Nuytsia : bulletin of the Western Australian Herbarium

ISSN 0085-441

NUYTSIA

Volume 4 Number 2
1983

WESTERN AUSTRALIAN HERBARIUM— DEPARTMENT OF AGRICULTURE

Cover

A representation of Nuytsia floribunda (Labill.) R.Br. ex Fenzl — the Western Australian
Christmas Tree. The journal is named after the plant, which in turn commemorates Pieter Nuijts, an
ambassador of the Dutch East India Company, who in 1627 accompanied the “Guide Zeepard” on
one of the first explorations along the south coast of Australia.

NUYTSIA

VOLUME 4 NUMBER 2
1983

WESTERN AUSTRALIAN HERBARIUM,
DEPARTMENT OF AGRICULTURE,
SOUTH PERTH, WESTERN AUSTRALIA

CONTENTS

A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Aus-
tralia. By Paul G. Wilson 135

Publication date of Volume 4 Number 1 262

Editorial Board

B. R. Maslin (Editor)
J. W. Green
P. G. Wilson

Editorial Assistant

J. W. Lee-Frampton

Western Australian Herbarium, George St, South Perth,
Western Australia 6151.

Nuytsia 4(2): 135-262 (1983)

135

A taxonomic revision of the tribe
Chenopodieae (Chenopodiaceae) in Australia

Paul G. Wilson

Western Australian Herbarium, George St, South Perth,
Western Australia 6151.

Abstract

Wilson, Paul G. A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia. Nuytsia
4(2): 135-262 (1983).

A taxonomic revision is presented for the tribe Chenopodieae (Chenopodiaceae) in Australia. Five genera
and fifty two species are recognised viz.: Chenopodium (24 species), Dysphania (10 species), Scleroblitum
(1 species), Einadia (6 species) and Rhagodia (11 species). One section, eight species, and seven subspecies
are described as new; six new specific and eight new infraspecific combinations are made. A number of
names are lectotypified, in some cases in a way different to that of earlier workers. The characters in the
fruit that have traditionally been used for circumscribing the genera are shown to be unsatisfactory; this
has led to the rearrangement of some species. The one species of Rhagodia described from outside the Aus-
tralian and New ^aland region is considered to be a species of Chenopodium.

Contents Page

Introduction 136

Morphology 137

Key to genera 138

Chenopodium 139

Subgenus Chenopodium 141

Sectio Chenopodium 141

Sectio Degenia 143

Sectio Leprophyllum 144

Sectio Desertorum 151

Sectio Auricoma 161

Sectio Rhagodioides 164

Subgens Ambrosia 166

Sectio Ambrina 166

Sectio Orthosporum 169

Dysphania 180

Scleroblitum 197

Einadia 199

Rhagodia 213

Doubtful or incorrect records 235

Excluded names 235

Acknowledgements 235

References 236

Figures 239

Maps 243

Index to scientific names 260

136

Nuytsia Vol. 4, No. 2 (1983)

Introduction

A historical survey of the treatment by different authors of the various genera now
included in the tribe Chenopodieae is given by Scott (1978a and b). He recognised
two subtribes, the Chenopodiinae and the Rhagodiinae, and distinguished them
principally on the presence of a dry membranous pericarp in the former and a fleshy
pericarp in the latter. With regard to the genera found in Australia, Chenopodium,
Dysphania, and Scleroblitum (as Chenopodium sect. Atriplicina) were placed in the
Chenopodiinae, while Einadia and Rhagodia were placed in the Rhagodiinae. I have
not found the nature of the pericarp to be a character suitable for discriminating
subtribes since, as detailed elsewhere, it can be succulent or dry in species which by
other criteria appear to be closely related. For this reason no subtribes are recognised
in this paper. Furthermore I have redefined the genera Chenopodium and Einadia so
as to include under both some species in which fruits are characteristically dry and
others in which they are succulent. Further notes on the history and circumscription
of some of the supraspecific categories are given under the respective genera or sec-
tions.

The species circumscription adopted here is rather broad (by the standards of
workers on the Chenopodiaceae in the European flora). Many taxonomic problems
remain, the resolution of which would often require extensive field work or the use of
experimental methods, neither of which are within the scope of the present study.
The names applied to the non-endemic species have been taken from recent papers
by workers in Europe, U.S.S.R., and North America and the relevant taxa are not
here critically revised.

The synonymy provided for the species endemic to Australia is as complete as I
have been able to ascertain. For other species the synonymy has been limited to
names based on Australian collections or to names used in significant publications on
Australian plants.

The author abbreviations used after botanical names follow those suggested in the
Draft Index of Author Abbreviations compiled at The Herbarium, Royal Botanic
Gardens, Kew (1980). The abbreviations for journal titles is generally in agreement
with G. H. M. Lawrence et al., Botanico-Periodicum-Huntianum (1968), while the
abbreviations used for books have, where possible, followed the suggestions of F. A. ^
Stafleu and R. S. Cowan, Taxonomic Literature, ed. 2 (1976-).

For each taxon usually two collections have been cited for each State in which it
occurs. However, all specimens studied have been determined and all specimens
bearing relatively precise geographical data have been mapped.

Paul G. Wilson, Tribe Chenopodieae

137

Morphology

Indumentum. A mealy indumentum is present to some extent in all species of
Einadia, Rhagodia, and Sclerobhtum, while it is also present in some species of
Chenopodium subg. Chenopodium. The mealiness consists initially of one-celled
spherical or irregularly shaped globules that may be white and dull, or glossy and
transparent. The spherical or irregularly shaped cells are supported on slender stalk
cells and both arise from the division of an epidermal cell (Esau 1977). Brian and
Cattlin (1968) imply a non-cellular morphology to those spherical globules that con-
stitute the mealy indumentum of Chenopodium album (and therefore, by inference,
other mealy members of Chenopodiaceae). This appears to be incorrect since other
workers (i.e. Wood 1923; Osmond et al. 1969, and Esau 1977) have described and
illustrated the origin of the indumentum from epidermal cells, while in some species
of Chenopodium a transitional series may be found ranging from the subsessile
globule to a slender multicelled trichome. In those cases where the mealy indumen-
tum is very dense the stalk cells are of different lengths so that the spherical cells are
arranged in layers; this is the situation in Chenopodium gaudichaudianum and
Rhagodia spinescens. The spherical cell usually collapses on drying as the organ on
which it has arisen matures. If the cell wall is relatively firm the spherical cell then
becomes saucer-shaped, as is common in many species of Chenopodium, Einadia,
and Rhagodia (e.g. C. desertorum, E. nutans and R. spinescens). If, however, the cell
wall is weak the cells may collapse to form a scurfy covering which is eventually shed
as irregular shaped flakes, or they may fuse together to form a silvery sheen on the
leaf or branch. The latter state is adopted in Rhagodia candolleana and R.
crassifolia, a circumstance which serves to distinguish these two from all other
species of the genus.

The mealy indumentum described above has been shown to have at least two
functions, but not necessarily in the same species. Wood (1923) demonstrated that
the collapsed cells in Atriplex vesicaria Benth. (which resemble those of Rhagodia
candolleana) assist in the prevention of water loss by the plant, while Osmond et al.
(1969) have shown that in some species of Atriplex the cells play an important part
in excreting salt. These two activities are probably of only minor significance in those
species of the Chenopodiaceae in which the mealy indumentum may be absent, or
sparse and soon lost. For instance in Rhagodia eremaea the globular cells do not fuse
together after collapsing but remain distinct as small scales which are shed as the
leaf matures. In this species, therefore, the mealy indumentum would appear to have
little influence on either the water or salt economies of the plants (see Hellmuth
1968, where referred to as R. baccata).

In Dysphania and in Chenopodium sect. Orthosporum are found both glandular
and simple, septate -acuminate hairs, characters which support the suggestion of a
close relationship between the two groups.

In Chenopodium sect. Ambrina both sessile glandular hairs and septate simple
slender hairs are found on the branches and leaves, while distinctive geniculate hairs
occur on the ovary.

138

Nuytsia Vol. 4, No. 2 (1983)

In Chenopodium Sect. Auricoma vesicular mealy hairs are found, characteristic of
those in section Leprophyllum. However, in addition, on the branches of the inflor-
escence occur contorted tubular, multi-cellular hairs that are evidently of a similar
nature to the vesicular since they grade into them.

Sexuality. The species in the tribe Chenopodieae considered here range in their sexu-
ality from dioecy through gynomonoecy to a monoclinous condition. In those taxa
that are dioecious the terminal flower of a compound cyme tends towards the bisex-
ual state; e.g. in a male plant of Rhagodia the terminal flower has a large pistillode,
and in a female plant more prominent staminodes, than do the lateral flowers. In
those taxa that are gynomonoecious the terminal flower of a cyme is bisexual while
the lateral flowers are female; this is typically the situation in Einadia. In Dysphania
the terminal flower is either bisexual or male (if the latter it possesses a prominent
pistillode), while the lateral flowers are male.

Pericarp. The pericarp varies from being diaphanous (in Dysphania), membranous
(in some species of Chenopodium), leathery (Chenopodium species), succulent
(Rhagodia etc.), or crustaceous (Scleroblitum). In some species (e.g. Chenopodium
desertorum) it can be either succulent or dry in the same subspecies, evidently de-
pendent to some extent on the environmental conditions. The unusual nature of the
crustaceous pericarp of Scleroblitum is described under that genus where it is noted
that although in this character it is unique among the Chenopodiaceae of Australia,
it is similar to Monolepis, a genus of small herbs found in Asia and America.

Key to genera

1. Plants shrubby, dioecious 5. Rhagodia

Plants herbaceous or weakly shrubby, monoecious or flowers polygamous. ... 2

2. Herbs with basal rosette of leaves; tepals 4, cartilaginous in fruit

3. Scleroblitum

Plants without a basal rosette 3

3. Plants glandular pubescent; tepals 1-4; stamens 1-2; embryo lateral or basal

2. Dysphania

Plants glandular pubescent or mealy; tepals (3) 5; embryo variously positioned .4

4. Tepals 5; stamens 1-3 in terminal bisexual flowers; seed horizontal; perennial

herbs (or weak shrubs) with mealy indumentum 4. Einadia

Tepals (3-)5; stamens 1-5; seed horizontal or vertical; mostly annual herbs
(rarely perennials or weak shrubs) with mealy or glandular indumentum or
glabrous Chenopodium

Paul G. Wilson, Tribe Chenopodieae

139

1. CHENOPODIUM L.

Linnaeus, Sp.Pl. 218 (1753). Lectotype: Chenopodium rubrum L. (see Britton and
Brown 1913).

Orthosporum (R. Br.) T. Nees, Gen. FI. German, ad t. [58] (1835). Chenopodium
sect. Orthospo'rum R. Br., Prod. 407 (1810). Lectotype: Chenopodium carinatum
R.Br. (see Ulbrich 1934).

Annual (rarely perennial) herbs, occasionally weak shrubs, mealy with minute
sessile vesicular hairs which may collapse to form a scaly indumentum, glandular or
glabrous. Leaves alternate, petiolate, chartaceous or somewhat fleshy, entire or vari-
ously serrate or dissected. Flowers small, bisexual or female, usually clustered. Peri-
anth 3 to 5 lobed, herbaceous or coriaceous, sometimes becoming cartilaginous in
fruit. Stamens (l-)5, free or united at base into a saucer-shaped disc. Style usually
very short; stigma 2 to 3(5). Pericarp membranous or rarely succulent. Seed lenticu-
lar to sub-globular, horizontal to vertical; testa crustaceous; embryo annular to hip-
pocrepiform; radicle centrifugal or (in vertical seeds) inferior. Seed shed either at-
tached to or free from perianth.

A cosmopolitan genus of over 70 species; 24 are recorded from Australia of which
14 are probably endemic.

Classification

The most recent outline of the infrageneric classification in Chenopodium is that
provided by Scott (1978b). In that paper he surveys the historical aspects and pro-
vides a considerable synonymy.

I have differed from Scott on certain points, principally in the typification of some
names and the circumscription or recognition of some sections. These matters are
discussed under the relevant taxa. The Chenopodium species here recognised as
being native or naturalized in Australia are classified as follows:

Chenopodium L.

subgenus Chenopodium

sectio Chenopodium, species no. 1
Degenia Aellen, species no. 2
Leprophyllum Dumort., species nos. 3-9
Desertorum Paul G. Wilson, species nos. 10-12
Auricoma Aellen, species nos. 13-14
Rhagodioides Benth., species no. 15
subgenus Ambrosia A. J. Scott

sectio Ambrina J. D. Hook., species nos. 16-17
Orthosporum R.Br., species nos. 18-24

140

Key to species

Nuytsia Vol. 4, No. 2 (1983)

1. Plants somewhat mealy at least when young, not glandular 2

Plants glandular pubescent, not mealy 18

2. Shrubs 3

Herbs (sometimes perennial) 6

3. Fruit dry 4

Fruit succulent 5

4. Spinescent shrub; leaves leathery, spathulate, to 3 cm long . 15. C. nitrariaceum

Unarmed soft-wooded shrub; leaves chartaceous, oblong-elliptic to hastate, to 4
(6) cm long 14. C. auricomum

5. Stigmas strongly penicillate 12. C. gaudichaudianum

Stigmas papillose 11. C. curvispicatum

6. Tepals 5, eventually free, almost orbicular, very sparsely mealy, not shed with

fruit; seed rounded on margin 13. C. auricomiforme

Tepals 3-5, if 5 then mealy and enclosing fruit when shed 7

7. Leaves variously toothed 8

Leaves entire or lobed 13

8. Seed deeply muricate 8. C. erosum

Seed smooth or minutely pitted 9

9. Perianth glabrous; seed horizontal or erect; perianth 3-5 lobed 10

Perianth mealy; seed horizontal; perianth 5-lobed 11

10. Perianth (at least of lateral flowers) shortly 3-lobed, urceolate

2. C. macrospermum

Perianth segments free 1. C. glaucum

11. Seed prominently keeled, finely pitted, dull; pericarp persistent . . 7. C. murale

Seed bluntly keeled, smooth to striate or weakly furrowed, pericarp readily de-
tached 12

12. Leaves considerably longer than broad, usually more than 3 cm long 3. C. album

Leaves about as long as broad, usually less than 3 cm long. . . .4. C. opulifolium

13. Plant foetid (smelling of rotten fish) 14

Plant not foetid 16

14. Tepals united to near apex; stamens 5 5. C. vulvaria

Tepals free except near base 15

15. Stamens 1 or 2 6. C. detestans

Stamens 5 lOd. C. desertorum subsp. virosum

16. Inflorescence a very slender panicle; tepals becoming black and enclosing fruit;

leaf lamina mostly c. 10 mm long; indumentum very thin 9. C. hubbardii

Inflorescence a narrow to broad panicle; tepals not black in fruit 17

17. Plant an erect annual; leaves mostly over 3 cm long with a very thin mealy indu-
mentum; panicle large, som.etimes 50 cm or more long 3. C. album

Plant prostrate to erect, usually a perennial; leaves always entire, usually less
than 2 cm long; panicle up to 5 cm long .10. C. desertorum

18. Stamens 1-2; seed lenticular, erect; leaves simple 19

Stamens 5; seed horizontal to erect; leaves simple or pinnatisect 25

19. Tepals 4 20

Tepals 5 21

20. Plant delicate, erect; flowers in open panicles (W.A.) 24. C. saxatile

Plant wiry, decumbent; flowers in dense axillary glomerules (New Zealand)

23. C. pusillum

21. Tepals strongly crested 22

Tepals rounded to acute on back or keeled 23

Paul G. Wilson, Tribe Chenopodieae

141

22. Perianth rostrate at apex; tepals free 21. C. cristatum

Perianth truncate at apex; tepals united 22. C. truncatum

23. Tepals keeled (at least towards apex) 19. C. carinatum

Tepals rounded on back 24

24. Tepals firmly united in lower half, completely covering fruit, usually black

20. C. melanocarpum

Tepals free to near base, not completely covering fruit, white ... .18. C. pumilio

25. Leaves serrate; seed horizontal to oblique 16. C. ambrosioides

Leaves pinnatisect; seed erect 17. C. multifidum

Subgenus Chenopodium

Sectio Chenopodium; A. J. Scott, Bot. Jahrb. Syst. 100:215 (1978).

Chenopodium sect. Pseudohlitum J. D. Hook in Benth. et J. D. Hook., Gen. PI. 3:52
(1880). Type: C. rubrum L.

Annual herbs more or less mealy. Flowers dimorphic in axillary and terminal pan-
icles. Perianth 3-, 4-, or 5-merous; tepals free or united in lower half. Stamens 1-5.
Seeds horizontal in terminal flowers, vertical in lateral flowers; embryo annular

Cosmopolitan with four species of which one is found in Australia.

1. Chenopodium glaucum L., Sp. PI. 1:220 (1753). Type: From Europe, n.v.

C. ambiguum R.Br., Prodr. 407 (1810). — C. glaucum var. ambiguum (R. Br.) J. D.
Hook., FI. Tasm. 1:313 (1857). — C. glaucum subsp. ambiguum (R.Br.) Murr et
Thellung ex Thellung, Mem. Soc. Natl. Sci. Nat. et Math. Cherbourg 38:196 (1912);
Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3/2:612 (1960). Type: Tasmania, R. Brown
(BM).

C. ambiguum var. majus Moq. in DC., Prodr. 13/2:67 (1849). No material cited.

C. ambiguum var. minus Moq. l.c — C. glaucum subsp. ambiguum f. minus (Moq.)
Aellen, Verb. Nat. Gesell. Basel 41:102 (1931). Type: Western Australia, Drummond
225 (iso:MEL).

C. littorale Moq., Chenop. Monogr. Enum. 24 (1840) nom. illeg. non (L.) Thunb.
(1815). Type: In nova Hollandia, Caley (holo:P).

C. glaucum var. littorale Rodway, Tasm. FI. 155 (1903). Type: No specimens cited.

C. glaucum L. var. paschale Fuentes, Bol. Mus. Nac. Chile 5:2 (1931) n.v., fide
Aellen, Feddes Repert. 26:47 (1929). Type: From Easter Island, n.v.

Annual herb, prostrate to decumbent or erect, to 0.5 m high, usually with several
angular branches arising from base. Leaves somewhat fleshy; lamina elliptical to del-
toid, 1-3 cm long, entire to sinuate dentate, often 3-lobed, obtuse, broadly cuneate at
base_ and passing into a petiole about half length of lamina, upper surface glabrous,
lower surface densely mealy. Inflorescence axillary and terminal of compact
glomerules arranged in narrow paniculate cymes. Flowers glabrous, dimorphic; ter-
minal flowers mostly hermaphrodite, tepals 3-4(5), free narrowly obovate, stamens 1-
4(5), stigmas subsessile, very short and thick, seed horizontal; lateral flowers female
or hermaphrodite, tepals 3(4), stamens 0-1, seed erect. Pericarp thin, free or loosely
attached to seed, somewhat green. Seed discoid, orbicular to oval, with rounded mar-
gin, c. 1 mm diameter; testa finely reticulate, smooth or minutely pitted, reddish
brown to black; embryo circular.

142 Nuytsia Vol. 4, No. 2 (1983)

NEW SOUTH WALES: Lake George, Apr. 1898, E. Betche (NSW); Front Beach,
L. A. S. Johnson 8248 (NSW).

VICTORIA: Thurla, A. C. Beauglehole 56212 (MEL); Wingan Inlet. N. G. Walsh 35
(MEL).

TASMANIA: Boomer, Mar. 1943, W. M. Curtis (HO); Stewarts Bay, J. H. Hemsley
6735 (HO).

SOUTH AUSTRALIA: Encounter Bay, 23 May 1933, J. B. Cleland (AD); 25 km S of
Naracoorte, D. Hunt 677 (AD).

WESTERN AUSTRALIA: Fitzgerald R., A. S. George 11274 (PERTH); Herdsman
Lake, K. F. Kenneally 1107 (PERTH).

Distribution. Southern temperate Australia. Cosmopolitan. Map 1.
Habitat. Usually in muddy eutrophic situations both inland and coastal.

Cbenopodium glaucum is a polymorphic species in which numerous infraspecific
taxa have been described. The ‘Australian’ plant is by some authors distinguished
from the European as a distinct species (C. ambiguum) and by others as subspecies
or variety of C. glaucum. Within Australia many variants are found, some of which
may be of very recent introduction while others could be indigenous. The typical
variant of C. ambiguum has deltoid 3-!obed leaves which are sinuate-dentate on the
margin whereas in the typical variant of C. glaucum the leaves are narrow-ovate to
narrow-elliptic with sinuate margins; Aellen (1960-1961) noted differences in the
flower morphology but this varies considerably between populations which have
similar leaf types. I consider it to be unprofitable to attempt to distinguish by name
any of the numerous variants found in Australia since to do so would require a
world-wide familiarity with the species and a knowledge of the application of the
many infraspecific names. Some of the variants found in Australia correspond closely
to those found in Europe so that subsp. glaucum also occurs in Australia if subsp.
ambiguum should be recognised as distinct. Most Australian collections appear to be
of a prostrate plant from boggy, often subsaline, situations; in Europe variants also
occur which have an erect habit.

Aellen (1960-1961) recognised C. glaucum subsp. ambiguum as being a native to
Australia, New Zealand, and Easter Island, and as a recent introduction to
Switzerland. The plant from Easter Island was earlier named C. glaucum var. pas-
chale by Fuentes.

Another rnember of section Cbenopodium (and the type of the genus) is C. rubrum
L. This species was recorded from Victoria by Ewart (1931), however, the specimens*
on which this record was based are C. glaucum; this error was pointed out by Willis

Railway Reserve, N.Melbourne, May 1910, C. French and J. A. Tovey (MEL); Coode Island, 23 March
1912, J. R. Tovey (MEL). J' ' n

Paul G. Wilson, Tribe Chenopodieae

143

Sectio Degenia Aellen, Magyar Bot. Lapok 25:56 (1927) n.v.; A. J. Scott, Bot.
Jahrb. Syst. 100:216 (1978b). Lectotype: Chenopodium macrospermum J. D. Hook.
(Scott op. cit.).

Annual herbs, sparsely mealy. Flowers slightly dimorphic in cymose clusters pan-
iculately arranged. Lateral flowers bisexual or female, perianth connate to near apex,
very shortly 3-lobed; stamens 0-3; stigmas short and thick; pericarp membranous,
free; seed usually vertical enclosed by sac-like perianth; embryo annular. Terminal
flowers bisexual and either similar to lateral flowers or perianth divided to near base
into 5 tepals; stamens 5; seed horizontal.

Three species in America and Europe. One species (C. macrospermum) introduced
into Australia.

2. Chenopodium macrospermum J. D. Hook., FI. Antarct. 341 (1846); Aellen,
Feddes Repert. 26:42 (1929). Type citation: ‘Falkland Islands; Berkely Sound and
St. Salvador Bay, near the sea; C. Darwin Esq., J. D. H.’ Type: n.v.

Erect annual to 1 m high branched from base. Leaves alternate, somewhat fleshy;
lamina rhombic or deltoid, c. 5 cm long, sinuate dentate, upper surface glabrous,
lower surface mealy when young, petiole slender ± equal to lamina. Inflorescence of
axillary sessile spike-like clusters forming terminal panicles. Flowers slightly dimor-
phic. Lateral flowers: perianth very shortly 3-lobed, urceolate, closely enclosing fruit,
c. 1.5 mm high, slightly succulent, glabrous, the lobes obscurely keeled; stamen soli-
tary; stigmas 2 very short and subsessile; pericarp thin, readily detached; seed erect,
flattened obovoid c. 1.5 mm long, reddish brown, glossy, testa faintly reticulate. Ter-
minal flowers either similar to lateral flowers but with perianth shortly 3-4 lobed, or
with perianth deeply 3-5 lobed and with 3-5 stamens.

WESTERN AUSTRALIA: Perry Lakes, 7.5 km west of Perth, K. F. Kenneally 7189
(PERTH); Lake Waneragup, 18 April 1978, B. K. Masters (PERTH).

Distribution. Western Australia, from Perth south to Bunbury. Probably a native of
North and South America and an adventive in Europe (Aellen 1929, Standley 1931).
Map 16.

Habitat. Peat on margin of slightly brackish lakes.

Chenopodium macrospermum is a very variable species. Aellen (1929) described
several infraspecific taxa, of these, the plant introduced into Western Australia cor-
responds most closely to C. macrospermum subsp. halophilum (Phil.) Aellen f.
subviride Aellen. For a comprehensive synonymy see Aellen and Just (1943). Cheno-
podium macrospermum subsp. halophilum was recorded for Australia by Probst
(1949 p. 63) but without the citation of collections. The earliest record known to me
is the one cited above dated 1978.

Aellen (1960-1961) described the terminal flowers of the clusters as being deeply 3-
5 lobed, having 3-5 stamens and horizontal to vertical seeds. In Western Australia
the terminal flowers are usually 3 lobed, rarely 4 lobed and only very occasionally
deeply 5 lobed, while the stamens are usually solitary (rarely 3) and the seed consist-
ently vertical.

144 Nuytsia Vol. 4, No. 2 (1983)

I am grateful to Pertti Uotila (H) for providing the identification of a Western
Australian collection of this species.

Sectio Leprophyllum Dumort., FI. Belg. 21 (1827); A. J. Scott, Bot. Jahrb. Syst.
100:217 (1978). Type: C. album L.

Chenopodium sect. Chenopodiastrum Moq. in DC., Prod. 13/2: 61 (1849) nom. illeg.
(superfluous name). Type: C. album L.

Chenopodium sect. Euchenopodium Ulbrich in Engler et Prantl, Nat. Pflanzenfam.
ed. 2, 16c:487 (1934) nom. illeg. Type: C. album L.

[Chenopodium sect. Chenopodium auct. pi. non sensu lectotypico.]

Annual herbs somewhat mealy. Flowers in cymose clusters spicate or paniculate.
Flowers hermaphrodite (and female), 5-merous, tepals united in lower half; stamens
5; stigmas 2-3. Pericarp membranous. Seed horizontal; embryo annular.

Numerous species mostly extra-Australian in origin, many of which are now
cosmopolitan weeds. Seven species are found in Australia.

3. Chenopodium album L., Sp. PL 1:219 (1753). Type: From Europe. Lectotype:
LINN 313.8, photo seen (see Brenan 1954 p.6).

C. lanceolatum R. Br., Prod. 407 (1810) nom. illeg. non Muehlenb. ex Willd.
(1809). — C. browneanum Roem. et Schult., Syst. Veg. 6:275 (1820). Type: Banks of
Paterson’s River, Oct. 1804, R. Brown (holo: BM).

C. probstii Aellen in Probst, Mitt. Naturf. Ges. Solothurn 20 (8):56 (1928); Aellen,
Verb. Nat. Ges. Basel 41:83 (1931). Type: Australia, N. Adelaide; Switzerland,
Derendingen in wool compost. See comments below. Syntype: Ward St., N. Adelaide,
16 May 1923, J. M. Black (AD 97810510).

Erect annual c. 1 cm high, usually much-branched, mealy white especially on
flowers and undersurface of leaves. Stem round to angular with green stripes. Leaves
thin, the lower petiolate, ovate rhombic, cuneate at base and variously dentate, the
upper narrowly ovate to lanceolate entire, more or less sessile, the longer ones c. 4-6
cm long. Inflorescence largely leafless of sessile flowers in cymose glomerules aggre-
gated in a large paniculate, spicate, or indeterminate arrangement. Flowers her-
maphrodite; tepals 5, united in lower half, broadly ovate, obtuse, membranous on
margin, weakly keeled (terminal flowers larger and more strongly keeled), green and
somewhat mealy. Stamens 5, glabrous, disc absent. Pericarp membranous, glabrous,
readily detached. Seed horizontal, lenticular, c. 1.2 mm diam., bluntly keeled; testa
black, glossy, smooth, striate or weakly and irregularly furrowed (reticulate in var.
reticulatum). Seed shed within the surrounding perianth.

QUEENSLAND: Tamborine Mt., Apr. 1947, M. S. Clemens (BRI); Charleville,
M. M. Biddulph 195 (BRI).

NEW SOUTH WALES: Cooma, 15 Mar. 1955, E. Gauba (CBG); Canberra, N. T.
Burbidge 7484 (CANB).

VICTORIA: SE of Wallacedale, A. C. Beauglehole 30512 (MEL); 3 mi [4.8 km] east
of Tawonga, R. V. Smith 64/67 (MEL).

TASMANIA: Flinders Is., J. Whinray 23 (MEL); Macquarie Plains, Mar. 1861,
W. M. Curtis (HO).

Paul G. Wilson, Tribe Chenopodieae

145

SOUTH AUSTRALIA: 5 km south of Blyth, A. E. Orchard 364 (AD); Adelaide,
B. Copley 1310 (AD).

WESTERN AUSTRALIA: Albany, G. Perry 259 (PERTH); Manjimup, R. D. Royce
1463 (PERTH).

NORTHERN TERRITORY: Harper Springs, 26 Jan. 1949, W. G. Steenson (NT).

Distribution. All Australian states and territories but rare in the tropics. Cosmopoli-
tan. Map 2.

Habitat. Widespread as a weed in agricultural areas.

The name C. album has here been used in a broad sense and may encompass sev-
eral segregate species recognised by some European workers, including C. suecicum
J. Murr, C. berlandieri Moq. and C. giganteum D. Don. No attempt is made here to
distinguish the numerous infraspecific taxa which have been described, since such an
undertaking would require the attention of a specialist who is familiar with the com-
plex from different parts of the world. This is beyond the scope of this paper.

A plant which is readily recognisable in fruit is Chenopodium album var.
reticulatum (Aellen) Uotila (1978 p. 31). This variety may be distinguished by the re-
ticulate testa; it occurs sporadically with the other variants across Australia.

The application of the name C. probstii is in doubt. The plant was originally as-
sumed by Aellen (1928 op. cit.) to have originated in South Australia and to have
been introduced into Europe as a contaminant of wool. Jorgensen (1973 p. 314),
while accepting it is a distinct species, throws doubt on its Australian origin and
suggests it comes from North America. It has never been listed in an American flora
and, if present in America, is presumably there included under C. album. In a study
of C. album and C. suecicum in northern Europe Uotila (1978) discusses C. probstii
and suggests that its taxonomy will only be understood after a thorough study has
been made of the biology of the species in the C. album complex of both North and
Central America. A lectotype should only be chosen after such a study is undertaken.

Aellen (1960-1961) and Beauge' (1974) suggest a possible synonyomy of C. probstii
with Chenopodium serotinum L. (which itself is considered to be a synonym of C.
album by Beaug6). Kjellmark (1934) determined the chromosome number as 2n=54,
i.e. as a hexaploid; it therefore corresponds to that of C. album and C. opulifolium,
q.v.

4. Chenopodium opulifolium Schrader ex Koch et Ziz, Cat. PI. Palat. 6
(1814); DC., FI. Francaise 6:372 (1815). — C. album subsp. opulifolium (Koch et Ziz)
Maire, FI. de I’Afrique du Nord 8:36 (1962). Type: From Palatinate (Pfalz),
Germany, Schrader (S photo seen), see nomenclatural note below.

?C. triangulare Forsskal, FI. Aegypt. Arab. 205 (1775). Type: From ‘Taaes’. Yemen,
n.v.

Very similar to (and sometimes considered conspecific with) C. album. It differs
from the latter species in having long-petiolate 3-lobed rhomboid leaves 2-3 cm long
and wide, strongly mealy below, and in having keeled strongly mealy tepals.

SOUTH AUSTRALIA: About Mortlock Experimental Station, Mintaro, 5 March
1969, D. E. Symon 6699 (ADW, CANB).

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Nuytsia Vol. 4, No. 2 (1983)

Distribution. Southern South Australia. Central and southern Europe, north Africa.
Habitat. A weed of agriculture.

This species is rather variable in morphology and difficult to distinguish from C.
album, another highly complex taxon. The only unequivocal record is the Symon
specimen cited above although other collections included by me under C. album may
belong to C. opulifolium. It was first recorded in Australia by J. M. Black (1912) who
cited a specimen from Henley Beach, South Australia but this was later seen by P.
Aellen (1927 in litt.) who considered it to be better placed under C. album. An ad-
ditional character useful for distinguishing the two species has been pointed out by
Beaug6 (1974 p. 305); he notes that in the seed of C. album the micropyle has the
form of a dimple (a scar) while in C. opulifolium it resembles a knob (i.e. it is con-
vex). I have been unable to confirm tbis observation in material of C. opulifolium
available to me.

Work by Cole (1961, 1962) and Wilson (1980) has shown that both C. album and
C. opulifolium are hexaploids (2n=54) unlike most other weedy species of the section
which are either diploids (2n=18) or tetraploids (2n=36). Battandier (1888-1890 p.
753) states that one can find all intermediates between the two species. It would
therefore be reasonable to accept a subspecies status for C. opulifolium as was done
by Maire (1962 op. cit.).

Nomenclature. In the publication by Koch and Ziz (1814 op. cit.) the name
‘Chenopodium opulifolium. Schrad in. lit.’ appears, without a description but with a
reference to ‘Ch. album Borkh. in Rhein Mag.’ i.e. to M. B. Borkhausen (ed.),
Rheinisches Magazin zur Erweiterung der Naturkunde vol. 1:115 (1793). Here there
is a description of the form of Chenopodium album found in the valley of the Rhine.
I have not attempted to lectotypify this species nor to confirm that the plant de-
scribed by Borkhausen is the same as that to which Schrader gave the name C.
opulifolium (although the description agrees). The name C. opulifolium was also
validly published by Decandolle (1815 op. cit.) with reference to the publication by
Koch et Ziz (1814), to ‘C. viride Loiseleur’ (1806) and to ‘C. erosum Bastarde’ (1814).
Beauge" (1974) cites as type a Schrader collection from ‘Palatinate’.

The name Chenopodium triangulare Forsskal (1775) was given by Beauge' (1974)
as a probable synonym of C. opulifolium-, it is based on a plant from Yemen and has
generally been placed in synonymy under C. murale L. (e.g. by Christensen 1922).
Beaug^ points out that the latter synonymy could not be correct and that the only
species found in Yemen to which the description applies is C. opulifolium. If this is
so then C. triangulare is the correct name for this species. The type was not seen by
Christensen and is not present in Copenhagen (herb. C); I have not attempted to
search elsewhere.

5. Chenopodium vulvaria L., Sp. PI. 220 (1753). — C. olidum Curtis, FI. Londinensis
lasc. 5, t. 20 (c. 1785). Type-. From Europe, n.v.

Spreading annual c. 20 cm high branching from base with a strong fish-like smell
(of trimethylamine). Branches soft, mealy with thin-walled vesicular hairs. Leaves
alternate; lamina ovate to broadly ovate or trullate, 1-2 cm long, entire, apex acute,
base cuneate, upper surface subglabrous, lower surface mealy, petiole slender c. half
length of lamina. Inflorescence of dense clusters of flowers forming short compact
axillary and terminal thyrses 1-2 cm long. Flowers hermaphrodite (terminal) and fe-

Paul G. Wilson, Tribe Chenopodieae

147

male (lateral); perianth subglobular c. 1 mm diameter, shortly 5 lobed, enlarging with
and enveloping the fruit, densely mealy outside. Stamens 5, glabrous, united at base
into a cup-shaped disc, completely absent in female flowers. Pericarp membranous,
papillose, somwhat adherent. Seed horizontal, lenticular with obvious keel, c. 1-2 mm
diameter; testa radially lineate.

NEW SOUTH WALES: Binnalong, Jan. 1922, J. J. Parry (NSW); Berridale, Nov.
1914, D. J. O’Rouke (NSW).

VICTORIA: Warrnambool, H. B. Williamson 1348 (MEL); Omeo, R. V. Smith 64/29
(MEL).

SOUTH AUSTRALIA: Bool Lagoon, D. Hunt 1938 (AD).

Distribution. Eastern New South Wales, Victoria, south-eastern South Australia.
North Temperate areas. Map 16.

Habitat. Principally a weed of horticulture.

Chenopodium vulvaria may be recognised by its fish-like odour and by its shortly
lobed perianth which encloses the fruit. Features which distinguish it from C. de-
testans are discussed under the latter species.

Chenopodium vulvaria was first recorded in Australia by Ewart (1908) who stated
that it was “introduced with impure agricultural seed, and now widely spread”; the
earliest dated collection I have seen from Australia is the one cited above made by
O’Rourke in 1914.

6. Chenopodium detestans Kirk, Trans, and Proc. N. Zeal. Inst. 9:550 (1877);
Murr, Allg. Bot. Zeitschrift 16:57 (1910); Allan, FI. N. Zeal. 1:228 (1961). Type ci-
tation: “Originally discovered by Mr J. D. Enys, who informs me that he has seen it
in several localities in Canterbury”. Lectotype: Trelissick Basin, Canterbury, New
Zealand, 4 Feb. 1876, T. Kirk 821 (holo: WELT; iso: WELT), (see Allan 1961).

Prostrate to ascending annual herb with foetid odour, numerous wiry branches to
15 cm long arising from base. Leaves alternate; lamina thin, trullate, 5-10 cm long,
entire or with a pair of lateral teeth, sparsely to moderately mealy with round vesicu-
lar hairs. Inflorescence of short dense axillary thyrses up to 5 mm long. Flowers
mostly hermaphrodite; perianth erect in young flower, becoming incurved and ob-
scuring fruit; tepals (4)5, thin, broadly ovate with short broad claw, cucullate and
fleshy at apex, united towards base, mealy outside. Stamens 1 or rarely 2, completely
absent in female flowers, divaricate at anthesis, ± equal to tepals; filament linear
glabrous; disc absent. Pericarp membranous, prominently papillose, somewhat ad-
herent. Seed discoid to lenticular with rounded margin, c. 1.2 mm diam.; testa punc-
tate, black.

NEW SOUTH WALES: Nimitybelle near Cooma, Dec. 1896, J. H. Maiden (NSW
143153).

Distribution. Near Cooma, New South Wales, probably introduced; South Island,
New Zealand.

Habitat. In New Zealand “lowland to montane open tussock-grassland and bare
places” fide Allan (1961).

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Nuytsia Vol. 4, No. 2 (1983)

Chenopodium detestans resembles C. vulvaria in having the foetid odour of rotting
fish caused by the chemical trimethylamine which is presumably located in the oil
glands that are plentiful in the leaves and in the cortex of the stems. Apart from the
different habit and larger leaves of C. vulvaria, it differs from C. detestans in having
almost completely united tepals, usually 5 stamens united into a disc at base, and an
obvious keel to the seed.

The orginal description of C. detestans and that provided by Allan (1961) stated
that the flowers have four stamens (a most unusual number) whereas in material I
have examined from New Zealand and Australia (including the lectotype) the
number in the hermaphrodite flowers has been one or rarely two.

Chenopodium detestans is presumably endemic to New Zealand (the solitary re-
cord from Australia, cited above, is probably an introduction), however, there is also
the possibility that it is an early introduction to New Zealand from South America
since it behaves as a weed and is quite distinct from other Australian or New Zealand
endemic species. The absence of recent Australian collections suggests that it is not
common in that country, although in the field it could have been overlooked or con-
fused with C. vulvaria.

Neither Kirk (1877 op. cit.) nor Allan (1961 op. cit.) suggested the probable affin-
ities of C. detestans and this was not mentioned by Aellen (1960-1961) or Scott
(1978b) in their conspectuses of the genus. Murr (1910) placed it in sect.
Chenopodiastrum (= sect. Leprophyllum), in which decision he was followed by
Maiden and Betche (1916). Cheeseman (1925) and Ulbrich (1934) suggested an affin-
ity to C. vulvaria, probably being influenced by the odour the two species have in
common; the latter author placed it with C. vulvaria in sect. Euchenopodium (= sect.
Leprophyllum). My studies suggest that Chenopodium detestans is most closely re-
lated to C. carnosulum Moq. (1849) and C. scabricaule Speg. (1902) both of which
are indigenous to South America (C. carnosulum is also recorded from Mexico and
Texas). In the broad sense the three taxa could reasonably be treated as being
conspecific for they are very similar in vegetative and floral characters. Aellen and
Just (1943) placed C. scabricaule in synonymy under C. carnosulum, however, Giusti
(1970) and Planchuelo (1975) considered them to be distinct being distinguished in
part by the free pericarp of the former species and adherent pericarp of the latter.
Chenopodium detestans has an adherent pericarp and thus in this character is more
similar to C. carnosulum', it differs from both C. carnosulum and C. scabricaule in
having a foetid smell, a character which is noticeably lacking (according to available
descriptions) from the two American species.

7. Chenopodium murale L., Sp. PI. 1:219 (1753). Type: From Europe, n.v. Figure
4B.

C. biforme Nees, PI. Preiss. 1:636 (1845). — C. murale var biforme (Nees) Moq. in
(1849). Type: ‘In arenosis prope Pointwater, Perth’, Preiss 1256

(iso: MEL).

C. congestum J. D. Hook., London J. Bot. 6:280 (1847). — Rhagodia congesta (J. D.
Hook.) Moq. in DC., Prodr. 13/2:51 (1849). — R. baccata var. congesta (J. D. Hook.) J.
P.‘,, Tasmaniae 312 (1857); A. J. Scott, Feddes Report. 89:7 (1978). — R.

Mlardieri var. congesta (J. D. Hook.) Benth., FI. Austral. 5:153 (1870). Type: ‘Hobart
Town,’ Oct. 1840, Gunn 868 (holo: K).

Paul G. Wilson, Tribe Chenopodieae

149

Erect branched annual to 1 m high, sparsely mealy with thin walled vesicular
hairs, sometimes foetid when crushed. Stem angular. Lower and middle leaves petio-
late, lamina thin, broadly triangular to ovate, 2-8 cm long, mealy when young, acute,
broadly cuneate at base, coarsely and irregularly dentate with slightly incurved
teeth. Inflorescence of compact cymes in rather open panicles, axillary and terminal.
Flowers hermaphrodite (or female); perianth sparsely mealy outside (becoming
glabrous with age); tepals 5, c. 1.5 mm long, united in lower third, broadly ovate, ob-
tuse, prominently keeled and slightly cucullate at apex, membranous and minutely
ciliate on margin. Pericarp prominently papillose, often brownish, persistent. Seed
horizontal, lenticular, 1-1.5 mm diam., prominently keeled; testa glossy, minutely
pitted; embryo circular. Seed shed with surrounding perianth.

QUEENSLAND; Brisbane, A. Dietrich 2307 (NSW); Rockhampton, P. O’Shanesy
210 (MEL).

NEW SOUTH WALES: Pimpara Ck., S. Jacobs 1089 (NSW); Lord Howe Is., A.
Rodd 1453 (NSW).

VICTORIA: Port Phillip, Apr. 1886, C. Walter (NSW); Port Fairy, 1894, H. B.
Williamson (MEL).

TASMANIA: Stanley, Feb. 1948, W. M. Curtis (HO); Hobart, L. Rodway 9398 (HO).

SOUTH AUSTRALIA: Innes Nat. Park, C. R. Alcock 4821 (ADW); Mt Serle HS., D
E. Symon 4025 (ADW).

WESTERN AUSTRALIA: Midstream, M. /. H. Brooker 2103 (PERTH); Port
Denison, R. A. Saffrey 1557 (PERTH).

NORTHERN TERRITORY: Alice Springs, 27 Jan. 1976, A. Mitchell (NT)-
Woodgreen HS, D. J. Nelson 2131 (NT).

Distribution. Principally extratropical Australia. Cosmopolitan. Probably indigenous
to the Mediterranean area and to southern and south-western Asia (Engstrand and
Gustafsson 1973). Map 5.

Habitat. A weed of agriculture and of disturbed areas, also coastal sands.

Numerous infraspecific taxa have been described from Europe but no attempt has
been made in this paper to relate these names to the plants found in Australia.

The name Chenopodium congestum (1847) was based on a collection found on the
south coast of Tasmania in which no fruits were observed by the author. Moquin
(1840), although he saw only the type collection, described the young fruit as being
slightly fleshy for which reason he transferred the species (but with reservations) to
Rhagodia. Later, J. D. Hooker (1857 op.cit.) while accepting it as a member of this
genus made it a variety of R. baccata in which decision he was followed by Bentham
(1870). Neither Rodway (1903) nor Curtis (1967) mention the taxon. Scott (1978a)
retained it as a variety of R. baccata although his description of the taxon applies to
R. candolleana s. str. and two of the four collections he cites belong to that species;
the other two I have not seen. It is curious that no mention was made by Scott of the
dentate leaves of the type which very obviously distinguish it from any species of
Rhagodia, while the nature of the pericarp and testa conclusively identify it as being

22493 — ( 2 )

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Nuytsia Vol. 4, No. 2 (1983)

C. murale. The young fruit of C. murale, when dry, has a leathery pericarp and it is
probably for this reason that Moquin considered it would be somewhat fleshy when
mature.

Chenopodium murale was presumably an early introduction to Australia. It may
be readily recognised by the usually glabrous leaves with prominent antrorse teeth
and by the morphology of the seed.

8. Chenopodium erosum R. Br., Prodr. 407 (1810); Moq., Chen. Mon. Enum. 33
(1840); Moq. in DC., Prodr. 13/2:68 (1849); J.D. Hook., FI. Tasm. 1:313 (1857). Type:
In collibus arenos: Insularum Kents Group in fretu Bass, Dec. 1804, R. Brown (holo:
BM; iso: BM). Figure 4A.

[C. murale auct. non L.: Benth., FI. Austral. 5:160 (1870) p.p. as to the R. Brown
specimen cited.]

“C. sp. aff. suecicum J. Murr”, Willis, Handb. PI. Viet. 2:86 (1973).

Erect slender or bushy annual to 1.5 m high. Stems ribbed, with red stripes,
glabrescent. Leaves papery; lamina triangular to ovate 3-15 cm long, 1.5-10 cm wide,
acuminate, base somewhat truncate, margin deeply incised, undersurface pubescent
with thin-walled bladder-hairs when young, becoming glabrous; petiole 1/3 to 1/2
length of lamina. Inflorescence of compact terminal and axillary paniculate cymes,
becoming open with, age. Flowers hermaphrodite and female, sessile; tepals 5, shortly
united at base, obovate, c. 1 mm long, with broad scarious margins and a fleshy mid-
rib, pubescent with thin-walled bladder-hairs when young, becoming glabrous;
stamens 5, glabrous, filaments linear, disc absent; ovary glabrous. Pericarp membra-
nous, dull, adherent to seed. Seed horizontal, lenticular with rounded margins, c. 1.5
mm diam.; testa black, glossy deeply muricate with radial striations. Seed eventually
falling free from perianth.

QUEENSLAND: Gayndah, 13 May 1917, C. T. White (BRI); Upper Logan, Anon.
284 (MEL).

NEW SOUTH WALES: Moona, Walcha, A. R. Crawford 464 (MEL); Coneac,
Johnson et Briggs (NSW 142705); Tinderry South Peak, J. H. Willis (MEL 80686).

VICTORIA: Bonang— Gelantipy Rd., A. C. Beauglehole 37698 (NSW); Gippsland,
F. Mueller (MEL 80689); 3 mi [4.8 km] NE of Suggan Buggan, J. H. Willis (MEL

TASMANIA: Kent Group, Bass Strait, Dec. 1804 R. Brown (BM).

SOUTH AUSTRALIA: South-west River, Kangaroo Is., J. B. Cleland (AD);
Ardrossan, J. G. O. Tapper 915 (AD).

Distribution. South-west Queensland to Bass Strait, southern South Australia; New
Zealand. Map 6.

Habitat. River banks, forest lands, often in disturbed areas.

This species is one of two in the sect. Leprophyllum apparently native to Aus-
tralia. Other members of the section which have been described from Australia are
now considered conspecific with European species and, therefore, probably represent

Paul G. Wilson, Tribe Chenopodieae

151

relatively recent introductions to that country. Chenopodium erosum itself has the
aspect of an introduced Eurasian species, however, I have been unable to match it
with any extra-Australian plant.

Although C. erosum is found over a wide area it has been collected only in-
frequently, Robert Brown’s collection of 1804 being the solitary one for Tasmania. In
New Zealand it is also widespread but rare; all collections in that country were made
this century (W. R. Sykes, CHR, pers. comm.).

9. Chenopodium hubbardii Aellen, Candollea 8:19 (1940); Aellen in Hegi, 111. FI.
Mitt.-Eur. ed. 2, 3:616 (1960). Lectotype: Mungallala, Queensland, 31 Dec. 1930,
C. E. Hubbard and C. W. Winders 6022 (holo: K n.v.; iso: BRI) lecto. nov.

Erect annual herb branching from base, c. 30 cm high. Branches slender, striate,
sparsely mealy when young. Leaves alternate; lamina thin, ovate to hastate with
rounded apex and lobes, c. 10(30) mm long, entire, upper surface sparsely mealy,
lower surface mealy grey due to the opaque discrete vesicular hairs; petiole slender
from half as long to as long as lamina. Inflorescence a terminal slender panicle c.
2 cm long (lengthening to 10 cm in fruit); flowers in compact glomerules (which be-
come separated with age) subtended by filiform bracts, the branches terminated by a
solitary bisexual flower (the others female). Flowers depressed globular c. 1 mm
diam.; tepals orbicular, sessile (or in fruit shortly clawed), densely mealy outside with
grey vesicular hairs. Bisexual flower, stamens 5, filaments linear broadened at base
and united into a thick disc; ovary glabrous; style very short (c. 0.1 mm); stigmas 2,
spreading, 0.3 mm long. Female flower: stamens and disc absent; ovary as in bisexual
flower. Fruiting perianth eventually black, depressed globular, hard, closely envelop-
ing fruit; tepals rounded on back, orbicular very shortly clawed. Pericarp brown to
black, finely papillose, adherent to testa. Seed lenticular, 1.5 mm diam. with rounded
margin; testa finely and irregularly pitted. Seed shed with enveloping perianth.

QUEENSLAND: 70 mi [112.6 km] S of Blackall, IF. J. Bissett E 233 (BRI); Morven,
S. T. Blake 10911 (BRI).

Distribution. South-central Queensland. A casual in Switzerland where introduced in
wool. Map 7.

Habitat. Usually recorded from heavy soils which become seasonally waterlogged.

Chenopodium hubbardii was considered by Aellen (1940 op.cit.) to be closely re-
lated to C. polygonoides and he later (1960-1961) included it in Chenopodium sect.
Polygonoidea. The two species are similar in appearance and both have persistent
pericarps and pitted testas but, whereas C. polygonoides has only a single stamen, C.
hubbardii has five. For this reason I am treating the latter as a member of section
Leprophyllum while C. polygonoides I am transferring to the genus Einadia. This
formal separation at the generic rank obscures the probable close relationship be-
tween the two species but the alternative, that of transferring all the species of
Einadia to Chenopodium, would cause considerable systematic confusion.

Sectio Desertorum Paul G. Wilson, sect. nov.

Herbae perennes vel suffructices, farinosi. Flores monoid; perianthium farinosum,
5-lobum. Stamina 5, discus patelliformis, sparse puberulus. Fructus baccatus vel

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Nuytsia Vol. 4, No. 2 (1983)

siccus; pericarpium glabrum, membranaceum vel succulentum, mesocarpio aquoso.
Semen horizontale, lenticulare vel depresso sphaericum; embryo annularis.

Typus: Chenopodium desertorum (J. Black) J. Black.

Perennial herbs or weak subshrubs largely covered with a mealy indumentum of
discrete globular vesicular hairs. Flowers monoecious, those terminal to the inflor-
escence and its branches male, the others female. Perianth 5 lobed, mealy outside
with vesicular hairs. Stamens 5, united at base into a sparsely puberulous disc. Fruit
baccate or dry; pericarp glabrous, free from seed, membranous to succulent with a
watery mesocarp. Seed horizontal, lenticular to depressed spherical; embryo annular.

Three species endemic to Australia.

The species in this section have, in floras and herbaria, frequently been confused
with each other, for they are similar in indumentum, leaf-shape, and in flower and
fruit morphology. In habit they are intermediate between herbs and shrubs being
either herbaceous perennials or weak shrubs with straggly branches. The fruits when
succulent have a watery interior and, on drying, leave either an inflated or a thin
wrinkled pericarp which is often sticky but lacks the viscous character of the meso-
carp in Rhagodia. Because of their often shrubby nature and their often succulent or
semi-succulent fruit, these species have sometimes either been placed in Rhagodia or
confused with one of its species. In that genus they are obviously misplaced; their
semi-woody habit, their monoecious flowers, and their watery fruits (when baccate)
clearly set them apart, while their habit and fruit also make them out of place in the
genus Chenopodium. To segregate this group of species at the generic level is unde-
sirable due to the similarity between some variants of C. desertorum and such north-
ern-hemisphere species of Chenopodium as C. vulvaria. I have therefore included
them in that genus, a course which is least disturbing of current taxonomic practice.
Within Chenopodium the species of section Desertorum have most in common with
those of subsection Lejosperma Aellen et Iljin of section Chenopodium (correctly of
sect. Leprophyllum) in which are included the species C. album, C. opulifolium, and
C. vulvaria.

10. Chenopodium desertorum (J. Black) J. Black, FI. S. Austral. 181 (1924) et in
Trans. Roy. Soc. S. Austral 49:272 (1925). — C. microphyllum var. desertorum J.
Black, Trans. Roy. Soc. S. Austral. 46:566 (1922). Type citation: ‘Murray lands; Port
Augusta westward to Ooldea’. Lectotype: Ooldea, Nov. 1921, D. Bates (holo: AD
97811131; iso: MEL 545186, NSW 143061) lecto. nov.

Rhagodia prostrata A. Cunn. ex Moq. in DC., Prod. 13/2:52 (1849) non C. prostratum
(Pallas) Roemer et Schultes (1820). Type citation: ‘In Nova-Hollandia (Cunning.l)’.
Lectotype: N.E. from L.R. [i.e. Lachlan River], 5 Aug., A. Cunningham (K) lecto.
nov.

C. microphyllum F. MuelL, Trans. Phil. Inst. Viet. 2:74 (1858) nom. illeg. non Thunb.
(1794). C. triandrum var. lanuginosum F. E. Haviland, Proc. Linn. Soc. New South
Wales 36:532 (1912) based on preceding. — C. pseudomicrophyllum Aellen, Candollea
8:8 (1939) based on C. microphyllum F. Muell. Lectotype: Bacchus Marsh, 1853, F.
Mueller (lower left-hand specimen of MEL 80671) (MEL) lecto. nov.

C. anidiophyllum Aellen, Candollea 8:9 (1939). Lectotype: Dubbo, Feb. 1886, E.
Eetche (holo: G n.v.; iso: NSW 143105) lecto. nov.

C. cochlearifolium Aellen, Candollea 8:10 (1939). Lectotype: Lower Loddon R., Vic-
toria, Nov. 1887, C. Walter (holo: G n.v.; iso: MEL 80675, NSW 142716) lecto. nov.

Paul G. Wilson, Tribe Chenopodieae

153

Annual or perennial herbs or subshrubs, prostrate to erect, branched from the
base, covered at least when young with vesicular hairs or (inflorescence) with
branched or simple tubular hairs. Leaves opposite or alternate, chartaceous to some-
what fleshy, subsessile or with slender petiole; lamina elliptic, deltate, hastate or or-
bicular, 5-20 mm long. Inflorescence a narrow to broad panicle, shorter than or much
exceeding terminal leaves, flowers clustered in sessile (basically 3-flowered) cymes;
terminal flower of inflorescence and of its principal branches male or bisexual, lateral
flowers female. Flowers sessile, depressed -globular, 1-2 mm diameter; tepals 5, ovate,
shortly united at base, thickened in centre, scarious along margins, densely vesicular
hairy outside, spreading at anthesis. Male flower: stamens 5; filaments linear, united
at base into a circular disc which is sparsely vesicular pubescent within; pistillode
usually minute. Female flower: staminodes absent; ovary glabrous; style 0.2-0.5 mm
long; stigmas slender. Fruit enveloped by perianth; pericarp membranous to slightly
succulent and then separated from seed by a watery sap, sometimes sticky. Seed
horizontal, lenticular with rounded margin, 1-1.5 mm diameter, testa almost smooth
to slightly rugulose or striate, black.

Key to subspecies

1. Floral indumentum of glistening irregularly shaped vesicular hairs

a. subsp. desertorum

Floral indumentum of dull, grey to white vesicular hairs 2

2. Plant prostrate to decumbent; leaves small to 10 mm long; flowers small,

c. 1 mm diameter in short inflorescences 3

Plant erect; leaves 5-20 mm long, flowers 1-2 mm diameter; inflor-
escences usually exceeding terminal leaves 4

3. Leaves elliptic to orbicular, often glabrescent above; plant without a

noticeable smell b. subsp. microphyllum

Leaves narrowly elliptic, mealy on both surfaces; plant with foetid odour

d. subsp. virosum

4. Branches slender, straight and rigid; leaves small (to 10 mm long), orbi-
cular to obovate, sparsely mealy; flowers small (c. 1 mm diam), sparsely to

moderately covered with sessile vesicular hairs e. subsp.rectum

Branches straight or flexuose; leaves various, densely mealy (at least below);
flowers with globular or branched often stipitate hairs . c. subsp. anidiophyllum

10a. subsp. desertorum

Erect, rounded, much-branched herb to 20 cm high. Leaves densely covered when
young with colourless, glistening, transparent, branched vesicular hairs; lamina del-
toid to orbicular, 3-8 mm long. Inflorescence congested or loose, narrowly pyramidal
or slender, exceeding the terminal leaves and often up to 3 cm long. Flowers 1.5-2
mm diameter; perianth with a felty indumentum of glistening transparent
irregularly shaped vesicular hairs. Seed c. 1.5 mm diameter.

NEW SOUTH WALES: 5 mi [8 km] W of Moulamein, C. W. E. Moore 3400
(CANB); 128 km E of Wilcannia, 14 May 1971, G. J. White (NSW).

VICTORIA: Annuello, A. C. Beauglehole 55816 (MEL); Wyperfeld Nat. Park A C
Beauglehole 29462 (MEL).

SOUTH AUSTRALIA: Tallaringa Well, T. R. N. Lothian 3836 (AD); 10 km W of
Blanchetown, D. J. E. Whibley 3741 (AD).

154 Nuytsia Vol, 4, No. 2 (1983)

WESTERN AUSTRALIA: Fitzgerald R. Nat. Park, K. Newbey 4142 (PERTH);
Salmon Gums, R. D. Royce 3539 (PERTH).

NORTHERN TERRITORY: Ormiston Gorge area, J. R. Maconochie 2244 (CANB).

Distribution. Northern South Australia south-eastwards to central and western New
South Wales and north-west Victoria; rare in southern Western Australia and the
Northern Territory. Map 6.

Habitat. Recorded from deep sand but evidently also found in other well-drained
soils.

The typical variant of this subspecies is a compact perennial with felty leaves and
flowers. In western Victoria a variant is found with a looser habit and large leaves
with a less dense indumentum; the hairs of this variant are of the same nature as
those of the type.

10b. subsp. microphyllum Paul G. Wilson, nom. et stat. nov. — Basic name C.
micTophyllum F. MuelL, Trans. Phil. Inst. Viet. 2:74 (1858) nom. illeg. non Thunb.
(1794). — C. triandrum var. lanuginosum F. E. Haviland, Proc. Linn. Soc. New South
Wales 36:532 (1912), based on preceding. — C. pseudomicrophyllum Aellen, Candollea
8:8 (1939), based on C. microphyllum F. Muell. Lectotype: Bacchus Marsh, 1853, F.
Mueller (lower left-hand specimen of MEL 80671) (holo: MEL) lecto. nov.

Rhagodia prostrata A. Gunn, ex Moq., op. cit.

C. cochlearifolium Aellen, op.cit.

[Rhagodia spinescens auct. non. R. Br.: Benth., FI. Austral. 5:155 (1870) p.p. as to A.
Cunningham specimen cited.]

[C. triandrum auct. non. G. Forster: Murr, Allg. Bot. Zeitschr. 16:56 (1910); Maiden
et Betche, Census N.S. Wales PI. 66 (1916).]

Intricately branched prostrate to decumbent perennial with prominent taproot;
not noticeably foetid. Leaves thin to slightly fleshy, elliptic to orbicular, shortly
petiolate, 5-10 mm long, undersurface covered with small simple rounded white vesi-
cular hairs, upper surface glabrescent. Inflorescence short, scarcely exceeding the ter-
minal leaves. Flowers c. 1 mm diameter; perianth covered with small round white
vesicular hairs. Seed c. 1 mm diameter.

NEW SOUTH WALES: 15 mi [24.1 km] W of Rankin Springs, J. He Nardi 945
(NSW); 12 mi [19.3 km] S of Mt. Hope, C. W. E. Moore 5945 (CANB).

VICTORIA: Nhill, G. D’Alton 26 (MEL); 15 mi [24.1 km] W of Shepparton, T. B.
Muir 2547 (MEL).

SOUTH AUSTRALIA: 5 km W of Mannum, D. G. Spooner 2737 (AD); Hincks Nat.
Park, D. E. Symon 6495 (ADW).

WESTERN AUSTRALIA: Lake Wagin, 1890, M. Cronin (MEL); 2 mi [3.2 km] NW
of Ongerup, K. Newbey 3632 (PERTH).

Distribution. Principally in south-eastern Australia from Eyre Peninsula eastwards;
also in south-west Western Australia. Map 8.

Paul G. Wilson, Tribe Chenopodieae

Habitat. Usually in heavy soil; often in woodland.

155

This subspecies is variable both in habit and leaf shape and I have been unable to
clearly separate the variants of which it is composed. The white spherical vesicular
hairs combined with the small relatively thin leaves and prostrate to decumbent
habit distinguish subsp. microphyllum from the other subspecies.

Typification. The lectotype of Rhagodia prostrata is » specimen in herb. K bearing
that name in Moquin’s handwriting; a fragment in herb. P (mounted with the holo-
type specimen of R. drummondii Moq.) is presumably an isolectotype.

The name R. prostrata was included by Bentham (1870) and Scott (1978a) as a
synonym of R. spinescens R. Br., an obvious misplacement since it is certainly a
synonym of C. microphyllum F. Muell. Although the name R. prostrata Moq. (1849)
is earlier than C. desertorum (J. Black) J. Black, it cannot be transferred at the rank
of species to the genus Chenopodium since there exists the validly published name C.
prostratum (Pall.) Roemer et Schultes (1820).

The varietal combination Chenopodium triandrum var. lanuginosum was evidently
published inadvertently by F. E. Haviland in 1912 as a new name based on C.
microphyllum F. Muell. It would appear that Haviland obtained the name from the
herbarium of the Botanic Gardens, Sydney, which had received it from J. Murr as
the manuscript name “forma lanuginosa” for a collection made by E. Betche in
February 1886 in Dubbo (NSW 143105) and a collection from the Wimmera in Vic-
toria (NSW 143082). The duplicates of these collections in herb. PAE (now in herb.
G) were later cited by Aellen (1939) as syntypes of C. anidiophyllum. The name C.
triandrum G. Forster was considered by Murr (1910) to be correct for the taxa here
included under C. desertorum; however, the former species belongs to the genus
Einadia although it bears strong superficial resemblances to members of the C. des-
ertorum complex. Haviland (and others) applied the name C. triandrum var.
lanuginosum to collections of C. desertorum subsp. anidiophyllum.

10c. subsp. anidiophyllum (Aellen) Paul G. Wilson, stat. nov. — C. anidiophyllum
Aellen, Candollea 8:9 (1939). Lectotype: Coolabah, March 1907, J. H. Maiden (G ex
herb. P. Aellen) lecto. nov.

[Rhagodia parabolica auct. non R.Br.: Benth., FI. Austral. 5:153 (1870) p.p. as to
Dalton specimen cited.]

Erect perennial herb or weak subshrub c. 20 cm high with slender branches.
Leaves thin; lamina ovate to broadly elliptic, deltoid or subreniform, 10-20 mm long,
mealy below with round vesicular hairs, glabrescent above (in typical variant); peti-
ole slender, 5-20 mm long. Inflorescence more or less equal to terminal leaves (typical
variant) to well exceeding them. Perianth c. 1.5 mm diameter covered with simple or
branched white vesicular hairs. Pericarp thin and close fitting or loose and somewhat
succulent when fresh (wrinkled and orange when dry). Seed c. 1.5 mm
diameter.

QUEENSLAND: Tambo, S. L. Everist 1455 (BRI); Enniskillen, C. T. White 11674
(BRI).

NEW SOUTH WALES: Condobolin, G. M. Cunningham 255 (NSW); Dubbo, Feb.
1886, E. Betche (NSW 143105, syntype).

156 Nuytsia Vol. 4, No. 2 (1983)

SOUTH AUSTRALIA: Tinga Tingana, Strezelecki Creek, D. E. Symon 5845 (ADW).

NORTHERN TERRITORY: 24 mi [38.6 km] S of Alice Springs, D. J. Nelson 666
(CANB).

Distribution. The typical variant is found in north-central New South Wales and
south-central Queensland; the “central Australian” variant is found in western New
South Wales, northern South Australia and the southern portion of the Northern
Territory. Map 9.

Habitat. Varies according to variant involved.

In the eastern limit of its distribution (near Dubbo) this subspecies is a weak per-
ennial with small ovate leaves (lamina to 10 mm long) and a thin close indumentum
on the flowers; towards the west the leaves increase in size (lamina to 15 mm long)
and become more orbicular, tbe plant is more erect, and the indumentum becomes
denser and thicker. There appear to be a number of local variants within the general
transitional dine and, in addition, plants may be found intermediate in character be-
tween subsp. anidiophyllum, subsp. microphyllum, and subsp. desertorum, particu-
larly in the areas of overlap of these subspecies.

In southern Queensland subsp. anidiophyllum frequently has a strong smell of
stale fish (as is also found in subsp. virosum) but this is not always the case as has
been noted by collectors. Several collectors also note that the plants are very palat-
able to stock. The distribution map for subsp. anidiophyllum indicates a marked dis-
junction between the central Australian and the eastern populations which is not
reflected in the morphology of the plants from the two areas; specimens collected
from north-eastern South Australia and north-western New South Wales are vir-
tually identical to those from the “Centre”, that is, they have an erect habit with
straight branches, and relatively large leaves and flowers, these have a thick vesicular
indumentum. Further east the plants become more straggly with thinner and smaller
leaves and flowers. The “central” Australian variant appears to be usually found on
deep sand while the “eastern” variant is generally found on clays or loamy soils.

I have included under subspecies anidiophyllum the central Australian plant
which has thicker and usually larger leaves than that found in central New South
Wales from where the type comes. Also included is a variant found in the northern
Flinders Range with rather thick reniform leaves. Although these and other regional
variants have features which give them a distinctive appearance there is
intergradation and there is considerable morphological variation. An additional
reason for not splitting up this complex further is the difficulty of expressing the
subtle differences in a manner which can be interpreted by others.

Some specimens have a habit and leaf shape intermediate between subsp.
anidiophyllum and Chenopodium curvispicatum, they also have a faint reticulate or-
namentation of the testa. It seems likely that these plants are the result of
introgressive hybridization between the two taxa, particularly since they are found in
areas where the putative parents also occur, that is in western New South Wales and
in the Flinders Range.

Some of the specimens referred to subsp. anidiophyllum are possibly
intermediates between the subspecies desertorum and microphyllum but this cannot
be recognised easily without field experience.

Paul G. Wilson, Tribe Chenopodieae

157

Typification. Paul Aellen cited three syntypes with his description of Chenopodium
anidiophyllum, these were as follows:

“Victoria: Wimmera; no date and collector; sent by F. v. Mueller as C. microphyllum
F.v.M. (Herb. Murr). New South Wales: Dubbo, 1886, E. Betche (Herb. Murr);
Coolabab, 1904, J. H. Maiden (Herb. Murr).” The specimens were in a collection of
chenopods sent in about 1909 to Josef Murr in Feldkirch, Austria by J. H. Maiden,
Director of the Royal Botanic Gardens, Sydney. They were identified by Murr as
Chenopodium triandrum Forst. f. (see Murr 1910). I have selected as lectotype the
Coolabah collection since it is the only syntype present in Paul Aellen’s herbarium
(now in herb. G). It is also an excellent representative of the plant generally referred
in herbaria to C. anidiophyllum. The Dubbo syntype cited above belongs to this
taxon but the syntype from Wimmera (NSW 143105) has greater affinity to subsp.
microphyllum.

Bentham (1870) included collections of this subspecies in Rhagodia parabolica
(which it somewhat resembles in leaf-shape) while Scott (1978a) cited collections of
subsp. anidiophyllum under both Rhagodia gaudichaudiana {- Chenopodium
gaudichaudianum) and R. deltophylla (= R. spinescens). While fragments of vegeta-
tive material of subsp. anidiophyllum and of the other species can be similar in ap-
pearance, the flowers are very distinct; when in fruit, the subspecies cannot be con-
fused.

lOd. subsp. virosum Paul G. Wilson, subsp. nov.

Herba perennis effusa foetida. Lamina fold ovata vel elliptica c. 10 mm longa,
cbartacea, acute vel obtusa, ad basim anguste cuneata. Inflorescentia anguste pyr-
amidalis, congesta; flores depresso globosi c. 1.5 mm diam.

Typus: 15 mi [24.1 km] N of Deniliquin, New South Wales, Oct. 1965, W. E. Mulham
S393a (holo: NSW).

Spreading prostrate to decumbent, perennial herb with numerous branches arising
from stock, smelling strongly of rotten fish (trimethylamine) when crushed. Root a
woody taproot. Branches slender closely mealy. Leaves opposite or alternate, closely
covered with grey vesicular hairs; lamina (ovate to) narrow-elliptic, c. 10 mm long,
chartaceous, apex acute to obtuse, base narrowly cuneate and passing into the short
petiole. Inflorescence a dense narrowly pyramidal terminal panicle not exceeding
leaves, composed of few-flowered clusters subtended by linear bracts c. 2 mm long.
Flowers sessile, depressed, c. 1.5 mm diam.; tepals oblong, obtuse, incurved, rounded
on back and cucullate at apex, densely vesicular pubescent outside, somewhat en-
larged in fruit. Terminal flower of inflorescence and of lateral clusters male or bisex-
ual; filaments glabrous passing at base into a thick circular disc sparsely vesicular
pubescent or glabrous. Lateral flowers female; staminodes and disc absent. Fruit en-
veloped by the incurved tepals; pericarp membranous or slightly succulent, smooth,
readily detached. Seed c. 1.5 mm diam., testa radially striate.

NEW SOUTH WALES: Morundah — Urana, C. W. E. Moore 1372 (CANB); 15 mi
[24.1 km] N of Deniliquin, W. E. Mulham S 393 (NSW); Wunnamurra, June 1920,
J. Sloane (NSW).

VICTORIA: Daylesford (?), 1878, R. Wallace 182 (MEL).

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Nuytsia Vol. 4, No. 2 (1983)

Distribution. South central New South Wales, possibly also in central Victoria.

Map 9.

Habitat. Clay soil.

The arrangement and morphology of the flowers of subsp. virosum are the same as
those found in the other subspecies of C. desertorum and it is with this species that
the plant’s affinities obviously lie. The subsp. virosum is rather similar to subsp.
micropbyllum in leaf shape although readily distinguished by its smell. This smell of
trimethylamine is also present in C. vulvaria and C. detestans; subsp. virosum may
be distinguished from C. vulvaria by the differently shaped leaves and perennial
habit and from C. detestans by the flowers which in that species have only one or
two stamens.

The single herbarium record of subsp. virosum from Victoria is accompanied by a
label in which the locality ‘Daylesford’ has been queried; until further collections be-
come available its occurrence in that State must remain in doubt.

The epithet ‘virosum’ is taken from the latin word virosus meaning foetid.

lOe. subsp. rectum Paul G. Wilson, subsp. nov.

Herba perennis erecta ad 40 cm alta cinereo-virida. Rami recti, divaricati, tenues.
Lamina folii late elliptica ad late obovata vel orbicularis, 5-10 mm longa, ad apicem
rotundata vel truncata. Inflorescentia tenuis; flores c. 1 mm diam.

Typus: Porcupine Hill, Musgrave Range, South Australia, 29 Oct. 1966, J. Z. Weber
180 (holo: AD; iso: PERTH).

Erect grey-green perennial herb or subshrub to 40 cm high. Branches straight, di-
varicate, slender, sparsely covered when young with small discrete rounded vesicular
hairs. Leaves alternate; lamina broadly elliptic to broadly obovate or orbicular, 5-10
mm long, when dry often leathery and bordered in red, apex rounded to truncate,
lower surface sparsely to moderately covered with small sessile vesicular hairs,
glabrescent above; petiole slender, half length of lamina. Inflorescence slender, equal
to terminal leaves; perianth c. 1 mm diameter, sparsely to moderately covered with
small grey round vesicular hairs. Fruit with membranous to slightly succulent peri-
carp. Seed 1 mm diameter; testa smooth.

NEW SOUTH WALES; 13 mi [20.9 km] SE of Mt Hope, C. R. Dunlop 1538 (CBG);
45 mi [72.4 km] NW of Balranald, W. E. Mulham W372 (NSW).

VICTORIA: Hattah Lakes Nat. Park, 20 Feb. 1969, G. Anderson (MEL); Nandaly,
H. B. Watts 376 (NSW).

SOUTH AUSTRALIA: 5 km S of Maralinga, B. Copley 2698 (AD); Hambidge Nat.
Park, T. R. N. Lothian 4270 (AD).

WESTERN AUSTRALIA: 15 mi [24.1 km] E of Zanthus, R. D. Royce 5577
(PERTH).

NORTHERN TERRITORY: Petermann Reserve, T. S. Henshall 764 (NT); Finke
River, Schwartz and Schultz 14 (MEL).

Paul G. Wilson, Tribe Chenopodieae

159

Distribution. Western New South Wales and north-western Victoria to Eyre Penin-
sula, the North West Plains of South Australia and the southern portion of Northern
Territory; also recorded from Zanthus in Western Australia. Map 8.

Habitat. Deep sand.

The subspecies rectum is distinctive in its erect habit and slender straight
branches. The leaves are usually more leathery when dry than in the other
subspecies and are often bordered by a reddish tinge.

In spite of the wide and disjunct distribution of subsp. rectum the plant is rather
uniform in habit and in flower size and morphology. At Maralinga in South Australia
where it has been collected by T. R. N. Lothian (No. 4022, AD), a plant intermediate
in indumentum between subsp. rectum and a Rhagodia species has also been found.
This plant has a more shrubby habit than subsp. rectum and has sterile or partially
sterile male flowers with a woolly disc; it appears likely that this plant (T.R.N.
Lothian 4023, AD) is a hybrid between subsp. rectum and a species of Rhagodia.
Also from Maralinga has been gathered a specimen (F. L. Hill 748, AD) which is
intermediate in appearance between subsp. rectum and subsp. desertorum (a plant
also found in this area). This ‘intermediate’ specimen is probably a hybrid between
the two subspecies.

The infraspecific epithet is taken from the Latin word rectus, straight, referring to
the straight spikes.

11. Chenopodium curvispicatum Paul G. Wilson, sp. nov. Figure 4D.

[R. spinescens R.Br. var. deltophylla F. Muell., Rep. PI. Babbage’s Exped. 19 (1859)
p.p. not as to lectotype. (See note below).]

Suffrutex debilis, dense farinaceus. Folia plerumque opposita; lamina deltoidea vel
hastata, obtusa, 1-1.5 cm longa et lata, dense farinacea. Inflorescentia paniculata,
cernua, 2-5 cm longa. Fiores globosi, extus dense farinacei, c. 2 mm diam.; peri-
anthium 5-partitum in fructu divaricatum, 5 mm diam. Flores terminales masculini;
stamina 5, glabra; discus patelliformis, sparse vesiculoso-pilosus; pistillodium
minutum. Flores laterales feminei, staminodia et discus absens, ovarium glabrum.
Fructus baccatus; pericarpium succulentum. Semen horizontale, lenticulare, margine
rotundato; testa prominente reticulata, nigra.

Typus: South Australia, 3 km E of Sanderston; growing on red-brown calcareous
loam; straggling subshrub c. 70 cm high; fruit red; 30 Mar. 1959, R. Schodde 1082
(holo; AD; iso: CANB, duplicates distributed to CAL, K).

Weak intricately branched spreading shrub c. 0.5-1 m high. Branches slender (the
ends frequently nutant), densely mealy when young with vesicular hairs. Leaves op-
posite or sub-opposite; lamina deltoid to hastate with obtuse to rounded angles, 1-1.5
cm long and wide; densely covered with an indumentum of white rounded vesicular
hairs; petiole slender, somewhat shorter than lamina. Flowers polygamo-monoecious,
sessile or shortly pedicellate in drooping pyramidal panicles 2-5 cm long; bracts lin-
ear, caducous. Male (or bisexual) flowers terminal to inflorescence and to its
branches; perianth 5-partite, globular, c. 2 mm diameter, densely covered outside
with white stipitate vesicular hairs, glabrous within; staminal filaments linear-terete,
united into a broad saucer-shaped disc which is sparsely vesicular hairy within; pis-
tillode globular, usually minute but sometimes equal to a normal ovary and fertile.

160

Nuytsia Vol. 4, No. 2 (1983)

with stigmas subulate, erect and usually minute. Female flowers found below male;
perianth similar to that of male flower, enveloping fruit until fully mature when it
hardens and opens out to 5 mm diameter, often becoming red above; staminodes ab-
sent; ovary globular, glabrous, style short with a pair of slender papillose stigmas.
Fruit a berry; pericarp succulent, filled with sap within, red (orange on drying). Seed
lenticular with rounded margin, c. 1.5 mm diameter; testa prominently reticulate
with a honeycomb matrix, black.

NEW SOUTH WALES: Balranald, 7 July 1971, G. M. Alehin (NSW); 30 mi
[48.3 km] W of Euston, T. R. Whaite 1838 (NSW).

VICTORIA: Hattah Lakes Nat. Park, 23 March 1969, G. Anderson (MEL); Raak
Salt Plain, A. C. Beauglehole 16089 (MEL).

SOUTH AUSTRALIA: 38 km N of Morgan, N. N. Donner 311 (AD); 22 km SW of
Renmark, Hj. Eichler 13742 (AD).

WESTERN AUSTRALIA: 20 km W of Naretha, D. W. Goodall 822 (PERTH); 8 mi
[12.9 km] S of Norseman, L. A. S. Johnson W182 (NSW).

Distribution. Southern Australia from south western New South Wales and north
western Victoria westwards to south eastern Western Australia. Map 10.

Habitat. Evidently in well-drained soils often associated with limestone.

This species has been confused in herbaria and literature with several other
species, principally Chenopodium gaudichaudianum, C. desertorum, and Rhagodia
spinescens. Mueller (1859) included specimens of C. curvispicatum (along with those
of R. spinescens) when he described R. spinescens var. deltophylla (cf. collection
“Desert of the Murray River”, Feb. 1847, F. Mueller, MEL 87962, a syntype of that
variety); Bentham (18/0) cited the Mueller collection and at least three other speci-
mens of C. curvispicatum under R. spinescens] J. M. Black (1948) included charac-
ters pertaining to C. curvispicatum under both R. spinescens var. deltophylla and C.
desertorum (as is also evident from his herbarium); while Scott (1978a) cited collec-
tions here referred to C. curvispicatum under both R. gaudichaudiana and R.
deltophylla.

Chenopodium desertorum may be readily distinguished from C. curvispicatum by
its smooth seed; it is also a small plant whereas C. curvispicatum is a straggly shrub.
Rhagodia spinescens is a dioecious shrub with stiff straight compact inflorescences, it
has a small perianth which never encloses the fruit and the berry is viscous when dry
Chenopodium gaudichaudianum, while similar to C. curvispicatum in habit, has a
broader more branched inflorescence; it differs most obviously in the stigma which is
markedly penicillate.

The specific epithet curvispicatum refers to the drooping spike-like panicles which
are typical of this species.

12. Chenopodium gaudichaudianum (Moq.) Paul G. Wilson, comb, nov.—
Rhagodia gaudichaudiana Moq., Chenop. Enum. 11 (1840). Type: Bale des Chiens
marins, Gaudichaud (holo: P n.v.; iso: BM). Figure 7F.

R. coralliocarpa S. Moore, J. Bot. 35:168 (1897). Type: Kilkenny Soak, June 1895, S
Moore (holo: BM).

Paul G. Wilson, Tribe Chenopodieae

161

Intricately branched scrambling shrub to 2 m high. Branchlets, leaves and inflor-
escence densely mealy with discrete cream-coloured vesicular hairs. Leaves alternate,
chartaceous (when dry), deltoid to narrow hastate with slender petiole 3-5 mm long,
in all 10-20 mm long. Inflorescence a pyramidal panicle 5-10 cm long with disjunct
clusters of flowers, the main axis and branches flexuose. Flowers polygamo-mon-
oecious, subsessile, those terminating the inflorescence or its branches male (with
small pistillode) or bisexual, the others female. Perianth 5-partite, depressed
globular c. 1.5 mm diam. at an thesis and 3.5 mm in fruit, eventually spreading and c.
7 mm diameter, persistent, densely covered outside with vesicular hairs. Bisexual
flowers: stamens 5, filaments linear-terete, united into a broad saucer-shaped disc
which is glabrous or vesicular puberulous; ovary depressed globular, style slender
0.5 mm long with a pair of delicate strongly penicillate stigmas c. 0.6 mm long. Fe-
male flowers without staminodes. Fruit a berry, at first enveloped by perianth but
which eventually becomes spreading; pericarp pale and membranous or orange to red
and succulent with sap filled interior (not viscous), eventually falling from perianth.
Seed lenticular c. 1.4 mm diameter with rounded margin; testa deeply reticulate with
a rough surface, dark reddish brown.

SOUTH AUSTRALIA: Yudnapinna HS., N. N. Donner 3205 (AD); Tarcoola, E. H.
Ising 1408 (AD); Maralinga, 21 Sept. 1960, H. Turner (AD).

WESTERN AUSTRALIA: Roebourne, Aug. 1884, F. S. Carey (MEL); Narrine, Sept.
1903, W. V. Fitzgerald^iNSW); 23 mi [37 km] S of Learmontb, A. S. George 1258
(PERTH); Miss Gibson Hill, SW of Warburton, A. S. George 4043 (PERTH).

Distribution. Karratha to Geraldton in Western Australia, south-east to the northern
edge of the Nullarbor Plain, eastwards to northern Eyre Peninsula. Map 1.

Habitat. Sand, clay or loam, often in slightly saline situations.

This species was recorded by Bentham (1870) and by Maiden (1916) as occurring
in New South Wales but the collections on which these records were based are of
other species, principally C. curvispicatum. Scott (1978a) recorded it from central
and eastern South Australia and from New South Wales, but the relevant collections
are either C. desertorum or C. curvispicatum. The strongly penicillate stigmas of C.
gaudichaudianum clearly distinguish it from all other species of Chenopodium and
all species of Rhagodia, while its scrambling habit with flexuose branches make it
readily recognisable in the field. It is uniform in habit and morphology throughout
its range and does not appear to hybridize with other species.

The male flowers of Chenopodium gaudichaudianum have a widely spreading peri-
anth at anthesis but this subsequently closes, while in female flowers the perianth re-
mains closed during anthesis, with only the stigmas protruding, then opens widely
under the mature fruit.

Sectio Auricoma Aellen, Feddes Repert. 69:69 (1964); Aellen in Hegi, 111. FI. Mitt.-
Eur. ed. 2, 3:577 (1960) nom. illeg. (without latin diagnosis). Type: C auricomum
Lindl.

Herbs or soft-wooded shrubs, mealy with vesicular hairs and (at least in the inflor-
escence) villous with tubular hairs. Leaves chartaceous. Flowers in compact
glomerules forming dense cylindrical spikes paniculately arranged, bisexual or fe-
male by abortion of stamens; tepals 5, united at first in lower half; stamens 5, fila-

Nuytsia Vol. 4, No. 2 (1983)

ments linear, disc absent; ovary glabrous, styles very short, stigmas 2 and slender.
Pericarp papery. Seed horizontal (rarely oblique or erect), lenticular; embryo annu-
lar.

Two species endemic to Australia.

This section is very similar to sect. Leprophyllum particularly with the addition to
sect. Auricoma of C. auricomiforme which forms a connecting link between them, as
has been noted by Murr et Thellung (1915). It differs from sect. Leprophyllum prin-
cipally in the presence of tubular hairs in the inflorescence.

13. Chenopodium auricomiforme Murr et Thellung, Viertiljahrsschr. Naturf.
Ges. Zurich 60:432 (1915); Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:615 (1960).
Type: ‘Kammgarnfabrik Derendingen bei Solothurn, auf Kompost von australischer
Schafvolde’, 1915, R. Probst (holo: G). Figure 4c.

[C. album auct. non L.: Benth., FI Austral. 5:160 (1870) p.p. as to Barton specimen
cited.]

Erect annual herb to 1 m high. Stem and branches rather thick, striped green and
yellow, sparsely mealy when young. Leaves chartaceous; lamina 3-nerved, narrowly
ovate to ovate, rarely hastate (upper ones lanceolate), entire or sparsely and coarsely
toothed, 2-10 cm long, weakly cuspidate, glabrescent above, mealy below when
young. Inflorescence a terminal panicle with slender densely flowered branches made
up of compact glomerules subtended by filiform bracts, axis villous with tubular
hairs. Flowers bisexual (or female by abortion of stamens), sessile or subsessile,
spherical, c. 1 mm diam.; perianth almost glabrous (a few vesicular and tubular
hairs), (4)5-lohed to middle or lower (often initially syntepalous becoming
choritepalous with age), lobes (or tepals) imbricate, broadly elliptic to almost circu-
lar, sometimes abruptly constricted at base into a short claw, yellow-green with
fleshy keel and scarious margins, ciliate; stamens 5; filaments linear, glabrous; disc
absent. Ovary glabrous; style very short; stigmas 2, slender. Fruit free from perianth;
pericarp membranous, papillate, pale brown, adherent. Seed horizontal, lenticular, c!
1 mm diameter, rounded on margin; testa minutely puncticulate.

QUEENSLAND: Minerva, S. T. Blake 7927 (BRI); Pittsworth, C. T. White 6659
(BRI).

WALES: Bow via Merriwa, July 1923, H. Sturt (NSW); Premer, June
1912, H. McMaster (NSW).

SWITZERLAND: Derendingen, 12 Aug. 1927, P. Aellen (NSW).

Distribution. South-eastern Queensland and north-eastern New South Wales- a cas-
ual in Europe. Map 11. ’

Habitat. Black soil plains and heavy loams, often in low-lying situations.

This species was first recognised by Murr and Thellung who discovered it in
Switzerland as an introduction with wool from Australia. Aellen (1960-1961) con
A to be also a native of New Zealand but it is not recorded by

Allan (1940 and 1961) as either native or introduced in that country nor is it present
in the D.S.I.R. herbarium in Christchurch, N.Z. (CHR). Murr and Thellung (1915)
corisidered it to be intermediate in morphology between C. album and C. auricomum
and subsequently Murr (1926) suggested it might be a hybrid between these species.

Paul G. Wilson, Tribe Chenopodieae

163

Both species were confused with C. auricomiforme by previous authors for Bentham
(1870) cited specimens under C. album and F. Mueller (1869 p. 11) under C.
auricomum. The earliest record is a pre 1848 collection by L. Leichhardt from
Liverpool Plains (MEL 88487). Chenopodium auricomiforme is clearly a distinct
species and shows very little variability throughout its range.

Chenopodium auricomiforme was placed by Aellen (1960) in section Polygonoidea
along with C. trigonon, C. stellulatum, C. polygonoides, and C. hubbardii. However,
it differs greatly from the first three species both in habit and in number of stamens.
It compares most closely to C. auricomum, particularly in hair and perianth charac-
ters, and I have therefore transferred it to sect. Auricoma.

14. Chenopodium auricomum Lindley in Mitchell, J. Exped. Int. Trop. Austral.
94 (1848); Moq. in DC., Prod. 13/2:460 (1849); Benth., FI. Austral. 5:159 (1870);
Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:616 (1960). Type: Collected along the
Narran R., 14 March 1846, T. L. Mitchell (holo: CGE n.v.; ?iso: MEL, NSW).

?C. auricomum f. subglabrum Aellen in Probst, Mitt. Naturf. Ges. Soloth. 20 (8):56
(1928); Aellen in Hegi, l.c. Type: Derendingen, Switzerland, n.v.

Soft-wooded bluish grey shrub to 2 m high. Branches slender, costate, thinly
mealy. Leaves alternate, oblong-elliptic to hastate, 2-4(6) cm long including a slen-
der petiole c. 1 cm, entire, chartaceous, apex and basal lobes rounded, upper surface
glabrescent, lower surface with a thin mealy indumentum of grey or very pale yellow
discrete globular vesicles. Inflorescence grey mealy to very pale yellow in a broad ter-
minal panicle c. 10 cm long with ovoid to cylindrical branches composed of dense 3-5
flowered glomerules subtended by small spathulate bracts; axis pubescent with
tubular hairs. Flowers bisexual, or female by abortion of stamens (sometimes with
minute staminodes), sessile, depressed globular, c. 1 mm diam.; perianth united in
lower 1/2 to 2/3, entirely pubescent outside (or only in distal half) with irregularly
shaped short-tubular and bladder hairs, lobes imbricate, broadly obovate, membra-
nous, ciliolate; stamens 5, filaments linear and neither thickened at base nor united
into a disc; ovary globular, glabrous, style very short, stigmas slender. Fruit envel-
oped by perianth and deciduous with it; pericarp chartaceous, free, white and
opaque. Seed horizontal (rarely oblique or erect), lenticular, c. 1.5 mm diam., margin
rounded; testa minutely reticulate.

QUEENSLAND: Cunnamulla, S. T. Blake 10629 (BRI); Noondoo Stn., S. L. Everist
814 (BRI).

NEW SOUTH WALES: Depot Glen, S. Jacobs 3076 (NSW); Mt King Stn., R. A.
Perry 5m (NSW).

SOUTH AUSTRALIA: 48 km NE of Billa Kalina Stn., M. Fagg 379 (AD);
Murnpeowie Stn., D. E. Symon 5653 (ADW).

WESTERN AUSTRALIA: DeGrey River district, N. T. Burbidge 1585 (PERTH):
Mt Augustus Stn., 8 July 1970, D. G. Wilcox (PERTH).

NORTHERN TERRITORY: Supplejack Stn., T. S. Henshall 2306 (NT); Mt Andado
Stn., T. S. Henshall 601 (NT).

Distribution. Western Queensland, central and western New South Wales, northern
South Australia, northern Western Australia, Northern Territory. A casual in
Europe. Map 11.

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Nuytsia Vol. 4 , No. 2 (1983)

Habitat. Clay soil and heavy loam in depressions and areas of periodic flooding.

This species was placed by both Bentham (1870 l.c.) and Ulbrich (1934) in the
same section as C. album, but Aellen (1960 l.c.) removed it to the monotypic section
Auricoma, in which decision he was followed by Scott (1978b). The shrubby nature
of C. auricomum as well as the presence of tubular hairs suggests that it should be
considered sectionally distinct from the annual species related to C. album.

Bentham’s (1870) suggestion that C. auricomum might be conspecific with C.
furfuraceum Moq. (1840) and C. acuminatum Willd. (1799) is unacceptable since
Chenopodium furfuraceum is a synonym of Rhagodia candolleana Moq. (1840) q.v.,
while C. acuminatum is an annual species with a sharply keeled margin to the seed,
it is native to Asia and not found in Australia.

Owing to its palatability to stock this species has virtually disappeared from large
areas of the pastoral country. It is also edible to humans and Max Koch (1895 in
sched.) records it as being eaten by Aborigines in western New South Wales.

Sectio Rhagodioides Benth., FI. Austral. 5:158 (1870); J. D. Hook, in Benth. et J.
D. Hook., Gen. PI. 3:52 (1880); Volkens in Engler et Prantl, Nat. Pflanzenfam. III.
la:61 (1892); Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c:489 (1934); A.
J. Scott, Bot. Jahrb. Syst. 100:214 (1978). Type: C. nitrariaceum (F. Muell.) Benth.

Spinescent shrubs with a thin mealy indumentum which eventually collapses and
coalesces to form a scaly silvery sheen. Flowers sessile in a spinescent panicle, uni-
sexual or bisexual; tepals 5 united towards base, enveloping mature fruit; ovary pu-
bescent; stamens 5, glabrous; disc absent. Pericarp membranous, pubescent. Seed
horizontal to vertical; embryo annular.

A monotypic section indigenous to Australia.

This section differs from sect. Auricoma principally in the spinescent habit, the
weak mealy indumentum which becomes scaly with age, and the pubescent pericarp,
characters which, individually, are indefinite in their application but which together
demarcate C. nitrariaceum as a very distinct taxon.

Scott (1978b) included in this section Rhagodia ulicina (Gand.) Paul G. Wilson (as
a species of Chenopodium), however, R. ulicina is a dioecious plant with a fleshy
pericarp and was originally placed in the genus Chenopodium only through
Gandoger’s ignorance of its morphology.

15. Chenopodium nitrariaceum (F. Muell.) F. Muell. ex Benth., FI. Austral.
5:158 (1870) ‘nitrariacea’; F. Muell., Icon. Austral. Salsol. PI. t. 28 (1890); S. J.
Oostroom et T. J. Reichgelt, Gorteria 1:21 (1962). — Rhagodia nitrariacea F. Muell.,
Trans. Phil. Inst. Viet. 2:72 (1858); F. Muell. Sec. Gen. Rep. 14 (1854) nomen.
Lectotype: Upper left-hand specimen on sheet bearing labels 1) ‘Ad ripam fluvii
Murray’, Feb. 1847, and 2) ‘Avoca’, 2 Dec. 1852, both leg. F. Mueller (MEL 80673),
see note below, lecto. nov.

C. lycioides Gand., Bull. Soc. Bot. France 66:224 (1919). Type: Warracknabeal, Vic-
toria, 8 Oct. 1903, F. M. Reader (holo: LY).

Paul G. Wilson, Tribe Chenopodieae

165

Divaricately branched spinescent shrub c. 1 m high. Branches slender, covered
with a grey metallic sheen due to fusion of collapsed vesicular hairs. Leaves alter-
nate, frequently clustered at nodes; lamina coriaceous, spathulate with rounded apex
passing at base into a short petiole, in all 10-30 mm long, sparsely mealy with minute
branched collapsed vesicular hairs. Inflorescence a terminal panicle with spinescent
branches to 15 cm long and wide or reduced to a single branchlet. Flowers subtended
by small linear to ovate bracts, clustered in irregular glomerules, unisexual and di-
oecious or bisexual. Perianth depressed globular to broadly turbinate, 1.5-2 mm di-
ameter; tepals 5, united in lower half, pubescent outside with small branched tubular
hairs. Male flowers; stamens 5; filaments ligulate, glabrous; disc absent; pistillode c. 1
mm long, with minute erect stigmas. Female flower: staminodes minute c. 0.1 mm
long or absent; ovary pubescent above with branched tubular hairs; style short or ab-
sent; stigmas 2, slender, to 1.5 mm long, pilosulose. Fruit enveloped by perianth, dry;
pericarp membranous, white, pubescent above, free. Seed horizontal to vertical, de-
pressed globular or concavo-convex, to sub-reniform, c. 1 mm diam.; testa punctate,
reddish-brown to black, glossy; embryo hippocrepiform to annular.

QUEENSLAND: Noondoo Stn., S. L. Everist 831 (BRI).

NEW SOUTH WALES: Bourke, May 1918, J. L. Boorman (NSW); 8 mi [12.9 km]
SW of Tulpa, C. W. E. Moore 5433 (CANB).

VICTORIA; Dimboola, 1947, E. T. Muir (MEL); Borung, 13 Nov. 1903, F. M. Reader
(MEL).

SOUTH AUSTRALIA: 3 km W of Hawker, B. Copley 218 (AD); 48 km W of
Tallaringa Well, T. R. N. Lothian 3847 (AD).

NORTHERN TERRITORY: 7 mi [11.3 km] E of Docker R. Settlement, P. K. Latz
968 (AD); 12 mi [19.3 km] SE of Tempe Downs HS., N. M. Henry 585 (NT).

Distribution. Central Australia south eastwards to north western Victoria, central
and western New South Wales, south central Queensland. A casual in Switzerland,
Germany, Holland and England. Map 15.

Habitat. Clay soil in seasonally waterlogged situations.

This species exhibits considerable variation in the shape of the flowers and in the
nature of the stamens and ovary. In some plants the flowers are either all male (with
a very small pistillode) or all female, in other plants the apparently male flowers
have well-developed pistillodes which sometimes form seed (but the stigmas remain
miriute), while in yet others the flowers are bisexual. The orientation of the seed also
varies. I have not been able to find characters by which the several variants that ap-
pear to be involved could be separated, partly because the herbarium material is
usually imperfect.

Typification. Mueller (1858) did not cite specimens with his description of Rhagodia
nitrariacea but he gave as its distribution ‘Throughout the interior of Australia, from
Arnhem’s Land to Lake Torrens, the Murray River and its tributaries’. The only
type material in herb. MEL is of four specimens mounted on one sheet (MEL 80673)
on which are two labels indicating that the specimens were collected along the Avoca
River on 2 December 1852 and along the Murray River in February 1847. I have
chosen the upper left-hand specimen as the lectotype but there is no indication as to
which label it relates. A syntype in herb. K was collected by Mueller along

22493 — ( 3 )

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Nuytsia Vol. 4, No. 2 (1983)

Sturts Creek, Western Australia, presumably in 1856. This had been identified as R.
nitrariacea by Mueller but was cited by Bentham (1870) under R. crassifolia; it is a
specimen of R. eremaea.

Chenopodium nitrariaceum bears a superficial resemblance to Rhagodia ulicina
but the two differ markedly in hair type and in flower and fruit characters.

Subgenus Ambrosia A. J. Scott, Bot. Jahrb. Syst. 100:211 (1978) Type: C.
ambrosioides L.

Sectio Ambrina J. D. Hook, in Benth. et J. D. Hook., Gen. PI. 3:51 (1880). Type: C.
multifidum L.

Ambrina Spach, Hist. Nat. Veg. 5:295 (1836). nom. illeg. (includes type of Roubieva).
Type: C. multifidum L.

Roubieva Moq., Ann. Sci. Nat. Bot. ser. 2,1:292 (1834). — Chenopodium sect.
Roubieva (Moq.) Asch. et Graebner, Syn. Mitteleur. FI. 5/1:21(1913). Type: C. multi-
fidum L.

Annual or perennial herbs with yellow glandular hairs, aromatic. Leaves
herbaceous, entire to pinnatifid. Flowers in dense axillary glomerules, bisexual and
female. Perianth united in lower half; stamens 3-5; stigmas 2-5. Pericarp membra-
nous, covered in part with geniculate glandular hairs. Seeds horizontal to vertical;
embryo hippocrepiform to sub-annular.

About 8 species from South and Central America some of which are now cosmo-
politan weeds. Two species are found in Australia.

The sect. Ambrina has been by some authors (e.g. Ascherson et Graebner 1913,
Aellen 1960-1961, Scott 1978b) lectotypified on the species C. ambrosioides. How-
ever, Spach (1836 op. cit.) when describing the genus Ambrina, included in it both C.
multifidum L. (as A. pinnatisecta Spach) and C. ambrosioides (among other species).
Moquin (1840) while accepting the genus placed C. ambrosioides in Ambrina sect.
Adenois (which he stated to be the typical section) and C. multifidum in the genus
Roubieva. The latter name was based on Roubieva Moq. (1834) with the one species
R. multifida (Chenopodium multifidum L.). The generic name Ambrina is therefore
illegitimate being superfluous and is to be typified by the type of the name which
should have been adopted, i.e. C. multifidum. J. D. Hooker (1880) clearly based the
sectional name Ambrina on Spach’s generic name and gave it the same broad cir-
cumscription. I consider that C. ambrosioides and C. multifidum belong to the same
section so the effect of a lectotypification different to that of other recent authors
(who have selected C. ambrosioides) is not significant.

I have accepted the name Chenopodium sect. Ambrina in the sense of J. D. Hooker
(1880). It was first used by J. D. Hooker in Flora Novae-Zelandiae 213 (1853) with-
out reference to an author or to a basionym. He included two species, C. carinatum
R.Br. and C. pumilio R.Br., and gave as a description ‘seed vertical’. The name if
treated as new is illegitimate, being superfluous since it includes the lectotype of the
section Orthosporum R.Br. (1810). Hooker probably intended it to be based on the
genus Ambrina and was following Moquin (1840) who included both C. carinatum

Paul G. Wilson, Tribe Chenopodieae

167

and C. pumilio in Ambrina sect. Adenois (the typical section of that genus). It is de-
batable whether Hooker’s two word description in his 1853 publication is acceptable
as a diagnosis, particularly since his description for the unnamed group, which con-
tained the rest of the species he placed in the genus, was ‘seed horizontal, rarely (in
C. glaucum sometimes) vertical.’ This being the case, and in order to retain current
nomenclature where possible, I am treating the 1853 publication of the name as
being invalid. Chenopodium sect. Ambrina therefore dates from 1880 and is based on
Ambrina Spach.

The species of section Ambrina are peculiar in having geniculate hairs on their
ovaries. This character, along with their distinctive smell, clearly distinguish them
from all other Chenopodium species found in Australia.

16. Chenopodium ambrosioides L., Sp. PI. 219 (1753). Type citation: ‘Habitat in
Mexico, Lusitania’. Type: n.v.

Perennial herb c. 1 m high with ascending branches, strongly aromatic, glabrous or
villous towards apex. Leaves herbaceous; lower ones elliptic to narrowly elliptic,
coarsely serrate or incised, narrowed at base into a short petiole, in all c. 10 cm long;
upper ones entire and smaller; under-surface prominently covered with sessile
globular amber-coloured oil-filled hairs. Flowers sessile in small glomerules forming
leafy or leafless panicles, sometimes spicate. Terminal flower of glomerule bisexual;
perianth 5-lobed to near base, glabrous, lobes cucullate and enclosing flower;
stamens 5, glabrous, filaments ligulate, anthers exserted; ovary pubescent in upper
half with shortly stipitate geniculate glandular hairs, style short with 3-4 slender
stigmas. Lateral flowers female; perianth depressed spherical shortly 5-toothed and
enclosing flower. Pericarp membranous, free, glandular pubescent above Seed hori-
zontal or oblique, obtusely lenticular, c. 1 mm diam., almost smooth, glossy; embryo
hippocrepiform.

QUEENSLAND: Moreton Island, L. Durrington 1464 (BRI); Carnarvon Gorge, T. J.
McDonald 215 (BRI).

NEW SOUTH WALES: Mt Harris Station, 1 May 1952, E. F. Constable (NSW);
Scarborough Park, Koyarah, E. F. Constable 5940 (NSW).

VICTORIA: Recorded for eastern Victoria by Willis (1973), no specimens seen.
SOUTH AUSTRALIA: Paradise, Adelaide, A. G. Spooner 3973 (AD).

WESTERN AUSTRALIA: Fremantle, P. G. Wilson 8687 (PERTH).

Distribution. Widespread in mainland Australia. Cosmopolitan, native of tropical
America. Map 10.

Habitat. Disturbed situations, often bordering streams or creeks.

I have not attempted to distinguish in this paper the various closely related species
recognised by Aellen (1960), nor have I considered the infraspecific taxa enumerated
by Aellen (1929) and by Aellen and Just (1943). Much of the Australian material
agrees well with C. anthelminticum L. as delimited by Aellen (C. ambrosioides var.
anthelminticum (L) A. Gray), and a few specimens from Rockhampton, Queensland,
correspond to Aellen’s description (1960-1961) of C. spathulatum Sieber (C.
ambrosioides f. spathulatum (Sieber) Aellen 1929) which he records from Sydney.

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Nuytsia Vol. 4, No. 2 (1983)

The former species is distinguished by its slender spike-like inflorescence branches
in which the bracts of the flower clusters are very small or absent, while the latter
species is recognised by its spathulate bracts. Comments on this species complex by
C. F. Reed in Flora of Texas (1969) and by J. A. Duke in Flora of Panama (1961) ap-
pear to be apposite to the situation in Australia; Duke remarks on C. ambrosioides:
‘polymorphically perplexing, this species, or some of its subspecies, varieties or
forms, is quite cosmopolitan’, while Reed, after commenting on two varieties, states
that: ‘combinations of all characters vary to a degree that separating them is very
difficult’.

Voroschilov (1942) and Aellen (1960-1961) recognised the following three taxa of
the C. ambrosioides complex as being present in Australia:

C. integrifolium Vorosch., J. Bot. URSS 27:42 (1942) n.v.; Aellen in Hegi, 111. FI.
Mitt.-Eur. ed.2, 3:596 (1960).

C. spathulatum Sieber ex Moq. in DC., Prod. 13/2:73 (1849); Aellen in Hegi, loc. cit.

C. suffruticosum Willd. subsp. remotum Vorosch. op. cit. n.v.; Aellen in Hegi, op. cit.
595.

I have not attempted to discriminate any of the above taxa.

This species is cultivated in some countries for its oil which is considered to have
anthelmintic properties while the leaves are used for making the drink known as
Mexican tea. The variety antbelminticum is also cultivated and its oil, as the varietal
epithet suggests, has a vermifugal effect, which is more pronounced than that of the
typical variety. An account of the oil content of the plant found in Australia is given
by Shapter (1941).

17. Chenopodium multifidum L., Sp. PI. 220 (1753). — Roubieva multifida (L)
Moq., Ann. Sci. Nat. Bot. ser. 2, 1:292 D834). — Ambrina pinnatisecta Spach, Hist.
Nat. Veg. 5:296 (1836). Type citation: ‘Habitat in Bonaria’ (Buenos Aires). Type: n.v.

C. multifidum f. spathulatum Aellen, Amer. Midi. Naturalist 30: 50 (1943). Type not
cited.

Perennial herb, prostrate, decumbent or erect, to 1 m high, much branched from
the base, aromatic. Branches sparsely to moderately covered with cottony hairs.
Leaves pinnatisect, to 6 x 3 cm; segments linear with recurved margins, under sur-
face densely covered with pale yellow glandular hairs and sparsely so with cottony
hairs. Flowers predominantly female, sessile in few flowered glomerules forming
spicate panicles. Bisexual flower: perianth hemispherical, shortly 5-lobed; stamens 5,
filaments ligulate, glabrous; ovary narrowly turbinate, with shortly stipitate amber-
coloured geniculate glandular hairs in upper half; stigmas 3-5, sessile, slender. Fe-
male flower: perianth obovoid (pouch-like), minutely lobed and enclosing pistil.
Fruiting perianth c. 2.5 mm long, coriaceous and reticulate-veined. Pericarp mem-
branous, tree. Seed erect, lenticular with rounded margin; testa reddish brown,
glossy, puncticulate; embryo hippocrepiform.

QUEENSLAND: Near Warwick, Nov. 1958, Shirley (BRI).

NEW SOUTH WALES: East Tamworth, R. H. Goode 226 (NSW); Hay, Feb. 1970.
B. M. Alehin (NSW).

Paul G. Wilson, Tribe Chenopodieae

169

VICTORIA: Seymour, R. V. Smith 64/105 (MEL); Coode Is., 23 Mar. 1912, J. R.
Tovey (MEL).

SOUTH AUSTRALIA: Pt. Adelaide, D. E. Symon 1233 (ADW); Semaphore, 19 June
1940, C. G. Samuel (ADW).

WESTERN AUSTRALIA: Bunbury, Sept. 1942, C. A. Gardner (PERTH); Harvey,
Dec. 1967, K. S. Cole (PERTH).

Distribution. Southern mainland Australia. Native of South America and now widely
distributed. Map 12.

Habitat. Disturbed areas; often in coastal sand.

The branches of some specimens have the upper leaves entire and spathulate and
it was evidently on this variant that the f. spathulatum Aellen (1943) is based.

As will be noted in the sectional synonymy this species is the type of the names
Chenopodium sect. Ambrina and Chenopodium sect. Roubieva. These two sections
have been considered distinct by some authors (e.g. Aellen 1960-1961, Scott 1978b)
with C. ambrosioides (L.) Spach accepted as the type of sect. Ambrina and C. multi-
fidum of sect. Roubieva. However, even if this typification is adopted, the two sec-
tions appear to be not deserving of separate status since sect. Roubieva is then only
distinguished by its erect seeds and pouch-like perianth, characters which appear to
be rather insignificant compared with those the two sections have in common, par-
ticularly the glandular leaves and the curious geniculate hairs on the ovary.

Sectio Orthosporum R.Br., Prod. 407 (1810); Benth., FI. Austral. 5:162 (1870); J. D.
Hook, in Benth. et J. D. Hook., Gen. PI. 3:51 (1880) see note 1 below; Volkens in
Engler et Prantl, Nat. Pflanzenfam. 3, la:61 (1892); Aellen, Verb. Naturf. Ges. Basel
44:308 (1933); Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed 2, 16c:494 (1934);
Aellen in Hegi, 111. FI. Mitt.-Eur. ed. 2, 3:576 (1960); Scott, Bot. Jahrb. Syst. 100:213
(1978). Lectotype: Chenopodium carinatum R.Br. (see Ulbrich 1934). See note 2
below.

Blitum sect. Orthosporum (R.Br.) C. Meyer in Ledeb., FI. Alt. 11 (1829); Moq. in
DC., Prod. 13/2:81 (1849). See note 3 below.

Orthosporum (R.Br.) C. Meyer ex T. F. L. Nees, Gen. PI. FI. German, ad t. [58]
(1835). See note 4 below.

Orthospermum (R.Br.) Opiz, Seznam rostlin Kveteny Ceske 70 (1852). See note 5
below.

Chenopodium sect. Ambrina J. D. Hook., FI. Nov.-Zel. 213 (1853) nom. invalid, non
J. D. Hook. (1880). Type; C. carinatum R.Br.

Chenopodium sect. Carinata Standley, N. Amer. FI. 21/1:11, 27 (1916) nom. illeg.
(superfluous name). Type: Chenopodium carinatum R.Br.

Note 1: J. D. Hooker (1880) and Volkens (1892) included Chenopodium tenue
Colla from Chile in the section Orthosporum. However, this species is conspecific
with C. multifidum L., a member of section Ambrina.

Note 2: A. J. Scott (1978b) nominated Chenopodium pumilio R.Br. as the
lectotype of section Orthosporum but since the choice of lectotype had already been
made by Ulbrich (1934) this earlier selection stands.

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Nuytsia Vol. 4, No. 2 (1983)

Note 3: C. A. Meyer (1829 op.cit.) described Blitum sect. Orthosparum without
reference to the name Chenopodium sect. Orthosporum R.Br. He did, however, in-
clude in his section the two species described by Brown under sect. Orthosporum and
therefore it can be assumed that the names are nomenclatural synonyms. Moquin
(1840) recognised Meyer’s section but excluded both the species described by Brown;
instead he placed them in Ambrina sect. Adenois. This error was corrected in a later
publication (Moquin 1849).

Note 4; The generic name Orthosporum (R.Br.) C. Meyer ex T. Nees (1835
op.cit.) was based on Blitum sect. Orthosporum (R.Br.) C. Meyer (1829 op.cit.) but
without any mention of Robert Brown or of the two species Brown had placed in
Chenopodium sect. Orthosporum. A. J. Scott (1978b p. 209) considered the generic
name Orthosporum to be illegitimate and gave as type O. rubrum (L.) T. Nees ( =
Chenopodium rubrum L.); this was one of the two species named and illustrated by
Nees under Orthosporum, the other being O. bonus-henricus (L.) T. Nees (= C.
bonus-benricus L.). However, since the generic name Orthosporum is indirectly
based on Chenopodium sect. Orthosporum R. Br. it has the same type which is C. ca-
rinatum R.Br. The name Orthosporum is therefore legitimate and correct should the
“carinatum group” be considered to warrant separate generic status.

Note 5: The generic name Orthospermum w'as attributed by Opiz (1852 l.c.) to
Robert Brown and he gave as a synonym “Orthosporum C. A. Meyer”. It would
therefore appear that the first name is an orthographic variant of the second, and
that it was based on Brown’s sectional epithet either directly, or indirectly through
Meyer’s combination under Blitum, or through the generic name Orthosporum C.
Meyer ex T. Nees.

Annual herbs, prostrate, ascending, or erect, pilose with segmented hairs and with
sessile or stipitate gland-tipped hairs, strongly aromatic. Flowers ebracteate, small in
dense axillary clusters (condensed thyrses) or paniculate (C. saxatile), bisexual and
feniale. Tepals 4-5, erect, united towards base, variously thickened and enlarged in
fruit; stamens 0-1(2) opposite anterior or (in 4-merous flowers) anterior-lateral sepal;
stigmas 2, subsessile, slender. Pericarp membranous. Seed erect, lenticular; funicle
very short; embryo hippocrepiform around apex of seed; radicle inferior. Fruit de-
ciduous with perianth, an abcission zone forming at apex of pedicel.

Distribution. Australia and New Zealand; almost cosmopolitan through recent
introduction. Seven species, six of which are endemic to Australia and one to New
Zealand.

The habit, hair types, and inflorescence structure of the species in this section are
similar to those found in Dysphania section Tetrasepalac, the principal differences
between these sections of the two genera being found in the seeds. In Dysphania sect.
Tetrasepalae the seed is globular or subglobular and the embryo either lateral or
basal. In Chenopodium sect. Orthosporum the seed is lenticular and the embryo
always hippocrepiform around its apex.

Hybridisation commonly occurs between the species of this section (apart from C.
pusillum and C. saxatile) and several names have been given to the crosses or
backcrosses; these are listed below with critical comments. Since the species them-
selves exhibit considerable variability the number of different variants is immense
and it is pointless to attempt to name them. Where possible, herbarium specimens of
apparetit hybrid origin have been designated by a formula, but since parentage is
often difficult to determine (unless a suite of specimens has been collected) often
only the more obvious parent has been indicated.

Paul G. Wilson, Tribe Chenopodieae

171

In addition to the complexities noted above it is apparent that intergradation is
present between at least some of the pairs of species, in particular, the following:

1. C. pumilio — C. carinatum

2. C. carinatum — C. melanocarpum

3. C. cristatum — C. melanocarpum

These are discussed below.

Names of hybrids — sect. Orthosporum

A. Chenopodium xbontei Aellen, Verb. Naturf. Ges. Basel 44:317 (1933).

Considered by Aellen to be a hybrid between C. carinatum and C. cristatum which,
with the hybrid, were found at Derendingen in Switzerland. Aellen divided it into
two varieties as follows:

nothovar. cristatiforme Aellen, op. cit. p. 318. Lectotype: Derendingen,
Switzerland, 9 Sept. 1923, R. Probst (G) lecto nov. Syntype: Luterbach, Switzerland,
16 Aug. 1933, R. Probst (G).

Stated by Aellen to be a hybrid between C. carinatum var. holopterum (i.e. var. ca-
rinatum) and C. cristatum. The two specimens cited above appear to have been de-
rived from C. cristatum and another species, possibly C. carinatum.

nothovar. submelanocarpum Aellen, op. cit. Lectotype Derendingen, Switzerland,
5 Sept. 1926, R. Probst (G) lecto nov. Syntype: Kettwig, Germany, 5 Oct. 1930, L.
Bonte (G).

Stated by Aellen to be a hybrid between C. carinatum var. melanocarpum (= C.
melanocarpum) and C. cristatum. The two specimens cited appear to have had such
an origin.

B. Chenopodium xchristii Aellen, op cit. p. 317.

Considered by Aellen to be a hybrid between C. carinatum var. holopterum (i.e.
var. carinatum) and C. pumilio. Aellen divided it into two varieties as follows:

nothovar. intermedium Aellen, op. cit. 317. Type: “Western Australia: 120 km
south of Perth, 1901, leg. Cecil Andrews, 1th collect, no. 697”. (holo: K n.v.).

The description and illustration provided by Aellen suggest that this is a variant of

C. pumilio.

nothovar. semiconnatum Aellen, op. cit. 317. Type: “In Nova Holandia ad portum
Jackson, 1830, leg. Gaudichaud” (holo: G n.v.).

The Gaudichaud specimen is one of the two syntypes of Ambrina carinata var.
parvifolia Moq. (1840). It is probably a variant of C. pumilio.

C. Chenopodium trigonocarpum Aellen, op. cit. 41:99 (1930). Lectotype:

Derendingen, Switzerland, 31 July 1929, R. Probst (G) lecto. nov.

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Nuytsia Vol. 4, No. 2 (1983)

In addition to the lectotype, Aellen cited one other specimen from Derendingen
(not seen) and stated that it was also known to him from New Zealand and Australia.
In a later paper Aellen (1933) placed C. trigonocarpum in synonymy under C. ca-
rinatum var. holopterum (= var. carinatum). The lectotype appears to be a hybrid
between C. melanocarpum and probably C. cristatum or between C. melanocarpum
and another hybrid involving C. cristatum.

18. Chenopodium pumilio R.Br., Prod. 40 (1810); Benth., FI. Austral. 5:163 (1870);
Aellen, Verb. Naturf. Ges. Basel 44:314 (1933).— Bbtum pumilio (R.Br.) C. A. Meyer
ex Steudel, Norn. Bot. ed. 2, 1:210 (1840); Moq. in DC., Prod. 13/2:82 (1849).—
Ambrina pumilio (R.Br.) Moq., Chenop. Monog. Enum. 42 (1840). Type: Kangaroo
Island, R. Brown (holo: BM n.v.).

Blitum glandulosum Moq. in DC., Prod. 13/2:82 (1849) sensu lectotypica fide Aellen

(1933).— Chenopodium glandulosum (Moq.) F. MuelL, Fragm. 7:11 (1859). C.

pumilio f. glandulosum (Moq.) Aellen, Verb. Naturf. Ges. Basel 44:315 (1933); Aellen
in Hegi, 111. FI. Mitt.-Eur. ed. 2,3:598 (1960). Type citation: “In Nova-Hollandia”
“(v.s. in h. Desf., Poir., DC. et Webb.)”. Type: n.v.

Chenopodium carinatum f. rubellum Aellen, Feddes Repert. 24:345 (1928). C.

pumilio f. rubellum (Aellen) Aellen, Verb. Naturf. Ges. Basel 44:315 (1933). Type ci-
tation: “Johannisburg (advent, aus Austr.) 1922, leg. C. E. Moss! no 6702 (Herb.
Pretoria).” Type: n.v.

Chenopodium x christii n-var. intermedium Aellen, Verb. Naturf. Ges. Basel 44:317

(1933). Type: 120 km S of Perth, Western Australia, C. Andrews 1st. Coll, no 697
(holo: K n.v.).

Chenopodium x christii n-var. semiconnatum Aellen, Verb. Naturf. Ges. Basel 44:317
(1933). Type citation: “In Nova Hollandia ad portum Jackson, 1830 lee
Gaudichaud'\ Type: G-DC n.v.

Chenopodium pumilio var. oblongifolium J. Black, FI. S. Austral, ed. 2, 289 (1948)
nom. illeg., sine desc. latina.

[Chenopodium carinatum auct. non R.Br.: F. MuelL, Icon. Austral Sals PI t 32

19??’29g a972)'p^T'^^^ ^

[Blitum carinatum auct. non (R.Br.) Steudek
252 f. 8-10 (1909).]

Reichenbach, Ic. FI. Germ. 24: 118, t.

Annual or short-lived perennial herb to 30 cm high and wide, usually with several
stems arising from base, aromatic. Stems slender, simple or branched, sparsely to
moderately pilose with both slender segmented and sessile to stipitate gland-tipped
hairs. Leaves herbaceous; lamina narrow to broad elliptic to ovate, 10-20 mm long
obtuse, entire to sinuate or obtusely lobed, sparsely pilosulose with simple and
gland-tipped hairs; petiole slender, c. half to equal length of lamina. Flowers in com-
pact axillary glomerules, subsessile or on slender pedicels up to 2 mm long, ovoid, c.
0.5 mm high; tepals 5, erect, naviculiform, united towards base, herbaceous, thin
sparsely pilosulose at apex or along midrib. Fruiting perianth usually crustaceous
and white, c. 1 inm high; tepals erect, narrowly naviculiform, rounded on back, con-
tiguous at apex but often disjunct in middle to expose fruit. Pericarp diaphanous

dull grey or translucent, sparsely papillose. Seed lenticular 1 mm long keeled or
rounded on margin.

§^78^.^^^^^^' Longreach, S. T. Blake 6615 (BRI); Bengalla, L. Leichhardt (MEL

173

Paul G. Wilson, Tribe Chenopodieae

NEW SOUTH WALES: 78 mi [125.5 km] W of Cobar, J. C. De Nardi 225 (NSW);
Broken Hill, P. L. Milthorpe 176 (NSW).

VICTORIA: Kul Kyne Forest, A. C. Beauglehole 40651 (MEL); Creswick, Whan 196
(MEL).

TASMANIA: Buckland, 24 Apr. 1912, R. A. Black (HO); Cape Barren Is., J. S.
Whinray 733 (AD).

SOUTH AUSTRALIA: Millswood, D. N. Kraehenbuehl 1878 (AD); Queenscliffe,
Kangaroo Is., J. G. 0. Tapper 1285 (MEL).

WESTERN AUSTRALIA: Wyola, T. E. H. Aplin 2203 (AD, PERTH); Pemberton,
M. Koch 2558 (PERTH).

NORTHERN TERRITORY: Burt Plain, G. Chippendale (NT 8033).

Distribution. Principally southern Australia; as an introduced weed in many parts of
the world and as an occasional weed in northern Australia. Map 13.

Habitat. Heavy loamy soils, sometimes slightly saline; frequently an agricultural
weed.

The name Chenopodium carinatum was used by Bentham (1870) to include ma-
terial correctly placed in C. pumilio. He was followed in this regard by Mueller
(1889-1891), J. M. Black (1924) and Ewart (1931). It was not until 1933, when Aellen
published a revision of the section Orthosporum, that the separate identity of both
species became widely accepted, although some Floras continue to unite them.

Chenopodium carinatum and C. pumilio are evidently closely related. It would ap-
pear that in Australia with the widespread dissemination of C. pumilio since the
coming of agriculture (in which industry it features as a prominent weed) it has
spread into areas previously only occupied by C. carinatum. Integradation between
the two is apparent from a study of herbarium material. The situation is compli-
cated by the considerable variation, found between different populations of C.
pumilio, in habit, leaf shape, and perianth structure. In spite of this variability I
have not recognised any infraspecific taxa since the variants do not appear to occupy
discrete areas and there appears to be a smooth gradation between them.

In Chenopodium pumilio, and possibly in other members of the section, the an-
terior tepal of hermaphrodite flowers subtends the solitary stamen and is initially
larger than the others. At anthesis the swollen base of the staminal filament becomes
turgid and causes the stamen and anterior tepal to spread widely. Later all tepals be-
come equal in size and assume an erect posture. The perianth usually becomes carti-
laginous and white in fruit but this change does not always occur and would appear
to depend on the conditions under which the plant is growing.

19. Chenopodium carinatum R.Br., Prod. 407 (1810); Benth., FI. Austral. 5:162
(1870) p. pte; Aellen, Verh. Naturf. Ges. Basel 44:312 (1933) exc. var. mel-
anocarpum. — Salsola carinata (R.Br.) Sprengel, Syst. Veg. 1:923 (1825). — Ambrina
carinata (R.Br.) Moq., Chenop. Enum. 41 (1840). — Blitum carinatum (R.Br.) C. A.
Meyer ex Steudel, Nom. Bot. ed. 2, 1:210 (1840); Moq. in DC., Prod. 13/2:82 (1849).
Type: Hawkesbury R., New South Wales, R. Brown (holo: BM).

174

Nuytsia Vol. 4, No. 2 (1983)

C. cristatum var. holopterum Thellung, Vierteljahrsschr. Naturf. Ges. Zurich 64:724
(1919). — C. holopterum (Thellung) Thellung et Aellen in Probst, Mitt. Naturf. Ges.
Soloth. 20(8):57 (1928). — C. carinatum var. holopterum (Thellung) Aellen, Verh.
Naturf. Ges. Basel 44:312 (1933). Type citation: ‘Mit dem Typus der Art und mit Ch.
carinatum, 1917, Probst. ’ Type: n.v.

Ambrina carinata var. parvifolia Moq., Chen. Mon. Enum. 42 (1840) pro parte as to
lectotype.— Pbtum glandulosum var. parvifolium (Moq.) Moq. in DC., Prod. 13/2:82
(1849). Lectotype: D’Urville (P, n.v.) see Aellen (1933).

C. carinatum f. foliosum Domin, Biblio. Bot. 22:65 (1921). Type: Queensland,
Brisbane River, A. Dietrich 968 (iso: NSW).

Annual herb, decumbent to erect, branching from the base, aromatic. Stems
simple or branched, 30 cm long, pilosulose with both slender and sessile to stipitate
gland-tipped hairs. Leaves herbaceous; lamina ovate to broadly ovate, 5-20 mm long,
shortly lobed to coarsely serrate, glandular puberulous and pilosulose below on veins;
petiole slender, rather shorter than lamina. Flowers subsessile, densely congested in
fruit; tepals 5, erect, united towards base, narrow-oblong, naviculiform, hirtellous on
keel. Fruiting perianth white, thickened and coriaceous; tepals prominently keeled
especially towards apex, truncate or acute, sparsely hirtellous on keel. Pericarp di-
aphanous, smooth. Seed lenticular c. 0.5 mm long, keeled over radicle and channelled
over cotyledons.

QUEENSLAND: Hughenden, S. T. Blake 6181 (BRI); Wandoon, C. E. Hubbard
5072 (BRI).

NEW SOUTH WALES: Kunghur, E. F. Constable 6578 (NSW); 1.5km N of
Wollombi, L. A. S. Johnson and B. G. Briggs 3287 (NSW).

VICTORIA: Suggan Buggan, A. C. Beauglehole 36646 (MEL); Near Deddick, 21 Jan.
1948, J. M. Bechervaise (MEL)

Distribution. Eastern Queensland and New South Wales, far eastern Victoria. Map
4.

Habitat. Frequently an agricultural weed or an invader of disturbed soils.

There has been much confusion over the application of the name Chenopodium
carinatum, this arose through Moquin (1840, 1849) not having seen the type of that
species or of C. pumilio and therefore including material of the latter under the for-
mer name. This confusion was continued by Bentham (1870) who, while recognising
both species placed most specimens of each under C. carinatum and included in it
the plant later recognised as the species C. melanocarpum. The situation was clari-
ried by Aell^ (1933) who, unfortunately, had an individual approach to the
international Code of Botanical Nomenclature and who, for his understanding of the
species, relied to a large extent on plants arising in Europe from seed introduced
there in wool.

While I recognise C. carinatum as a distinct species, hybrids with C. pumilio, C.
melanospermum, and C. cristatum are common, probably largely through the move-
ment of seed of these species due to agriculture and commerce; this is particularly
apparent in Europe where numerous hybrids have been documented and in some
cases named. The species exhibits considerable geographic variation in leaf and peri-
anth form m particular in the tepals which range from being slightly to strongly

Paul G. Wilson, Tribe Chenopodieae

175

keeled. When strongly keeled the fruiting perianth has a form approaching that of C.
melanocarpum. Specimens with a particularly prominently keeled and hardened
perianth from central and South Australia and from north-western Western Aus-
tralia have, ill herbaria, sometimes been placed in C. carinatum and sometimes in C.
melanocarpum. I have placed them with the latter taxon, as did Aellen who named
this variant f. leucocarpum, although I recognise that the plants with this form of
fruit have evolved in several independent lines: some have affinity to C. carinatum,
some (a portion of the western element) probably to C. melanocarpum, while others
which 1 have segregated under f. leucocarpum may represent a hybrid between C. ca-
rinatum and C. melanocarpum.

20. Chenopodium melanocarpum (J. Black) J. Black, Trans. Roy. Soc. S. Austral.
58:173 (1934). — C. carinatum var. melanocarpum J. Black, Trans. Roy. Soc. S. Aus-
tral 46:566 (1922); Aellen, Verb. Naturf. Ges. Basel 44:313 (1933); Aellen in Hegi,
111. FI. Mitt.-Eur. ed. 2, 3:599 (1960). Lectotype: Broken Hill, New South Wales,
Sept. 1918, per T. G. B. Osborn (AD 96732197) lecto. nov.

[C. carinatum auct. non R.Br.: Benth., FI. Austral. 5:162 (1870) p.p. as to Drummond
165.]

[C. holopterum auct. non (Thell.) Thell. et Aellen: J. Black, FI. S. Austral. 683 (1929)
“C. holocarpum Thell. et Aellen”. See note below.]

Annual herb, prostrate, with numerous stems arising from the base, aromatic.
Stems simple, slender, to 30 cm long, pilosulose with both slender segmented and
sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy; lamina broadly
elliptic c. 15 mm long, bluntly lobed to almost entire, glandular puberulous and
pilosulose below along veins; petiole slender about half length of lamina. Flowers
subsessile to shortly pedicellate -(densely congested in fruit); tepals 5, erect, united in
lower half, narrow-oblong, keeled, hirtellous; stigmas 2, subsessile, slender. Fruiting
perianth bluntly stellate in cross-section, crustaceous, black (or pale fawn in f.
leucocarpum, firmly united in lower half and completely covering fruit; tepals
strongly and bluntly keeled, sparsely hirtellous on keel. Seed lenticular, 0.5 mm long,
keeled on margin over radicle, channelled over cotyledons; pericarp diaphanous, dull,
smooth or minutely puberulous towards apex, somewhat adherent to seed.

This species comprises two forms which may be distinguished by the colour of
their fruiting perianths.

20a. forma melanocarpum
Fruiting perianth hlack.

QUEENSLAND: 35 mi [56.3 km] SW of Eulo, M. law 43 (BRI); Boatman Station,
S. L. Everist 2792 (BRI).

NEW SOUTH WALES: Mt. Derriwong, G. M. Cunningham 774 (NSW); 12 mi
[19.3 km] E of Turlee, J. De Nardi 340 (NSW).

SOUTH AUSTRALIA: Mann Range, 22 Aug. 1954, J. B. Cleland (AD);
Amoorinyirra Hill, D. J. E. Whibley 1124 (AD).

WESTERN AUSTRALIA: 80 mi [128.7 km] E of Carnarvon, C. A. Gardner 6051
(PERTH); Mileura Stn., P. G. Wilson 8987 (PERTH).

176 Nuytsia Vol. 4, No. 2 (1983)

NORTHERN TERRITORY: Palm Valley, A. C. Beauglehole 10403 (PERTH); 4 mi
[6.4 km] N of Alice Springs, R. Swinbourne 563 (NT).

Distribution. Arid and semi-arid areas of Australia south of 22° latitude. Map 7.

Habitat. Well-drained situations either in sand or on sides of rocky hills.

The synonymy cited above is indicative of the difficulty experienced by different
workers in fitting a nomenclatural framework onto a variable and freely
interbreeding group. Bentham (1870 op.cit.) placed material of C. melanocarpum
(and of C. pumilio) under C. carinatum, while Black (1922 op.cit.) still recognising it
as part of that species, gave it varietal status, in which decision he was followed by
Aellen (1933, 1960-1961). Later Black (1934 op.cit.) raised var. melanocarpum to the
rank of species and in this he has been followed by workers in most Australian
herbaria. The course adopted by Black is preferable since the typical form of C. mel-
anocarpum differs appreciably in perianth and leaf shape from C. carinatum.

The spelling “C. holocarpum” was inadvertently used by J. M. Black (1924) in-
stead of C. holopterum. This error was later noted by Black in Trans. Roy. Soc. S.
Austral. 58:174 (1934) when he published the name C. melanocarpum as a new com-
bination.

20b. forma leucocarpum (Aellen) Paul G. Wilson, comb, nov.— C. carinatum var.
melanocarpum f. leucocarpum Allen, Verb. Naturf. Ges. Basel 44:313 (1933).
Neotype, here chosen: Ashburton River, Minderoo, Oct. 1905, A. Morrison (holo-
PERTH; iso: K). . v •

Fruiting perianth pale fawn in colour.

Distribution. North western Western Australia (Port Hedland-Exmouth Gulf area),
central and South Australia, eastern Queensland, New South Wales and Victoria’
Map 4.

Habitat. As for the typical form.

No collections were cited by Aellen for this forma, however, a duplicate at herb K
of the Morrison specimen which has been designated as a neotype has been deter-
mined by Aellen as f. leucocarpum.

21. Chenopodium cristatum (F. Muell.) F. Muell., Fragm. 7:11 (1864); Benth., FI.
Austral. 5:163 (1870); F. Muell., Icon. Austral. Salsol. PI. 31 (1890); Aellen, Verb.
Naturf. Ges. Basel 44:310 (1933).— Blitum cristatum F. Muell., Trans. Phil. Inst.
Viet. 2:73 (1858). Lectotype. Raro in planitiis sterilioribus subsalsis prope montis
Flindersianos, Oct. 1851, F. Mueller (holo: MEL 19998) iso: PERTH) lecto. nov.

Annual herb, prostrate to ascending with several stems arising from base, aro-
matic Stems slender, mainly simple, pilosulose with both slender segmented and
sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy; lamina elliptic
to broadly elliptic, entire to coarsely toothed, sparsely puberulous. Flowers
subsessde, tepals 5, erect, acuminate, strongly keeled and hirsute, ciliate, united in
lower half. FruAing perianth spherical, c. 2 mm diameter, strongly 5-winged, white
(rarely grey to black), tepals cartilaginous, strongly crested (vertically winged), semi-

Paul G. Wilson, Tribe Chenopodieae 17^

orbicular, rostrate, completely enclosing the fruit. Pericarp diaphanous dull, papil-
late towards apex or all over. Seed lenticular, c. 0.5 mm long, the margin slightly
keeled over radicle and grooved over cotyledons.

QUEENSLAND: Oakwood Stn., L. S. Smith and S. L. Everist 846 (BRI);
Thargomindah, Apr. 1885, Spencer (MEL).

NEW SOUTH WALES: “Birdwood” NE of Pooncarie, R. P. Green 177 (NSW); W.
of Stoney Crossing, D. L. W. Henderson 327 (NSW).

VICTORIA: 23 km NNW of Manangatang P.O., A. C. Beauglehole 55812 (MEL); nr.
Lakes Ranfurly and Hawthorn, May 1937, J. H. Willis (MEL).

SOUTH AUSTRALIA: Koonamore, Hj. Eichler 12492 (AD); “Markaranka”,
Morgan, D. E. Symon 3557 (ADW).

WESTERN AUSTRALIA: Esperance, R. A. Rose 3005 (PERTH); W side Fraser
Ra., P. G. Wilson 5898 (PERTH)

NORTHERN TERRITORY: Trephina Gorge, B. G. Briggs 1229 (NSW); 2 mi
[3.2 km] S of Weepita Bore, G. Chippendale (NT 2499).

Distribution. Arid and semi-arid regions of Australia south of 20° latitude. Map 14.
Habitat. Frequently in red sandy loam, often long creek lines.

This species may be readily recognised when in fruit by the strongly crested ros-
trate tepals. It is most similar to C. truncatum which may be distinguished by the
firmly united tepals and the absence of a rostrate apex to the perianth.

As is noted in the species description, the fruiting perianth while normally white is
sometimes grey to black in colour. In some cases these melanized plants have obvi-
ously arisen through hybridisation with C. melanocarpum (i.e. when the grey fruited
form has been collected with the two parents); in other cases the fruit shape is
typical of C. cristatum while the colour is black and no plants of C. melanocarpum
have been observed in the area.

22. Chenopodium truncatum Paul G. Wilson, sp. nov.

Herba prostrata vel adscendens cum pilis simplicibus et pilis glandulosis
subsessilibus pilosula, caulis numerosis ex caudice exorientibus. Folia modice
carnosa; lamina elliptica obtusa c. 10 mm longa, Integra vel leviter undulata,
glanduloso puberula. Flores in glomerulis axillaribus depositi; tepala 5 versus apicem
connata, c. 0.5 mm longa; stamina 0-1, glabra. Perianthium fructiferum
cartilagineum, late obtriangulare, c. 2 mm latum, apice applanato, marginis dentatis
vel breviter laceratis, tepalis firme connatis. Pericarpium diaphanum, nitidum,
glabrum. Semen erectum, lenticulare, c. 0.7 mm longum, margine rotundato.

Typus: 65 mi [104.6 km] W of Henbury HS on road to Kings Canyon, 10 Dec. 1968
P. K. Latz 255 (holo: AD; iso: CBG, NT).

Annual or (short-lived) perennial herb, prostrate or ascending, with numerous
stems arising from rootstock, aromatic. Stems slender, pilosulose with both slender
segmented and sessile to shortly stipitate gland-tipped hairs. Leaves slightly fleshy;

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Nuytsia Vol. 4, No. 2 (1983)

lamina elliptic c. 10 mm long, obtuse, entire or slightly undulate, glandular
puberulous all over and pilosulose below along veins; petiole slender about half
length of lamina. Inflorescence of discrete axillary glomerules. Flowers ebracteate,
subsessile, bisexual and female (the terminal always bisexual), ovoid, c. 0.5 mm high;
tepals 5, united to near apex, hirtellous on midrib; stamen 0-1, glabrous, anther
shortly exserted. Fruiting perianth cartilaginous, broadly obtriangular in plain view,
flat above (or the lobes slightly ascending), lateral margins toothed or shortly
lacerate, shortly hirtellous, white or black, c. 2 mm wide, the tepals firmly united to
apex. Pericarp diaphanous, glossy, glabrous. Seed erect, lenticular c. 0.7 mm high,
margin rounded (neither grooved nor keeled).

QUEENSLAND: Thylungra, S. L. Everist 5717 (CANB); Oakham Stn., Windorah,
28 Aug. 1959, R. S. Stranger (BRI).

NEW SOUTH WALES: Near Mt Murchison, Jan. 1887, R. H. Kennedy (MEL);
Calindry Stn., J. H. Leigh W183 (NSW).

SOUTH AUSTRALIA: NE corner of Lake Gairdner, B. Lay 341 (AD); 61 km E of
Dalhousie Springs, T. R. N. Lothian 1917 (AD); 10 km W of Quinyambi HS, D. J. E.
Whitley 5557 (AD).

NORTHERN TERRITORY: Old Andada Homestead, A. C. Beauglehole 27967
(NT); 5 mi [8 km] SE of Harper Springs Homestead, G. Chippendale (NT 6468).

Distribution. South-west Queensland, eastern New South Wales, northern and
central South Australia, the southern half of the Northern Territory. Map 3.

Habitat. Usually associated with deep red sand.

Specimens of this species have, in Australian herbaria, generally been placed with
Chenopodium cristatum, in some cases with the suggestion that they may be hybrids
with C. melanocarpum. Chenopodium truncatum differs from C. cristatum most ob-
viously in having a fruiting perianth which is truncate above (not with a rostrate
apex), the tepals firmly united to apex (not free) and not so markedly crested; the
pericarp is glabrous and glossy (not papillate and dull) and the lenticular seed has a
rounded margin (not keeled or grooved). From C. melanocarpum it differs in having
almost entire leaves (not strongly lobed) and in having crested tepals. Seven of the
collections (representing about 15 per cent of the total) consisted of mixed material;
in five C. cristatum was also present, in one C. melanocarpum, and in the other C.
pumilio. This suggested that C. truncatum was of hybrid origin. However, most
sheets of this species contained more than one specimen and in no case was there any
evidence of introgression. In addition hybrids between C. cristatum and C. mel-
anocarpum which I have observed on sheets of mixed material have a different ap-
pearance, being intermediate in morphology in both leaf and fruit characters.

The illustration provided by Aellen (1933) for the fruiting perianth of C. xbontei
var. suhmelanocarpum resembles the fruiting perianth of C. truncatum (except for
the union of the tepals in the latter). I have examined the two syntypes of var.
submelanocarpum and find them to be intermediate between C. melanocarpum and
C. cristatum as was stated by Aellen.

The specific epithet refers to the flat upper surface of the fruiting perianth.

Paul G. Wilson, Tribe Chenopodieae 179

23. Chenopodium pusillum J. D. Hook., Handb. N. Zeal. FI. 231 (1864);
Cheeseman, Man. N. Zeal. FI. ed. 2, 407 (1925); Allan, FI. N. Zeal. 1:228 (1961). Type
citation: ‘Sandy shores oF Lake Taupo’. Type: Colenso (K, n.v.).

[C. pumilio auct. non R.Br.: J. D. Hook., FI. Nov.-Zel. 1:214 (1853).]

Small decumbent annual with wiry stems radiating from base, c. 10 cm diam., pi-
lose with simple segmented hairs and with sessile or subsessile gland-tipped hairs,
aromatic. Leaves somewhat fleshy; lamina suborbicular to broadly ovate, obtuse, c.
5 mm long, entire or sparsely and bluntly dentate, sparsely puberulous above,
glandular puberulous below; petiole c. half length of lamina. Flowers in compact
axillary glomerules, shortly pedicellate; tepals 4, erect, c. 0.5 mm long, membranous
apart from the narrowly oblong midrib, united in lower half, pilose, eventually en-
larging to 1 mm long but not obscuring fruit. Stamen 1. Pericarp membranous, free
from seed. Seed compressed obovoid to lenticular, 0.5-0.8mm long; testa glossy,
finely puncticulate, dark reddish brown.

NEW ZEALAND: Christchurch, gravel ballast, 23 Feb. 1949, A. J. Healy (CHR);
Lake Lyndon, T. Kirk 1079 (CHR); Marlborough, on formed road, 23 Apr. 1952, L.

B. Moore (CHR).

Distribution. New Zealand.

Habitat. ‘In sandy and open places’ (Allan 1961); frequently a coloniser of disturbed
sites.

This species has been confused with Chenopodium pumilio which it resembles.
The two may be distinguished by the much smaller and more orbicular leaves of C.
pusillum, the 4-merous perianth, and by the membranous margin to its lobes which
do not become hardened in fruit. Chenopodium pumilio is also found in New
Zealand (presumably as an introduction). When growing in the same environment as

C. pusillum, the two are very similar in habit and general appearance.

24. Chenopodium saxatile Paul G. Wilson, sp. nov.

Herba annua erecta subtilis aromatica ad 30 cm alta, pilis debilibus glanduliferis et
pilis simplicibus rigidis modice pilosa. Caulis et rami tenues. Folia alterna; lamina
chartacea, ovata vel elliptica 10-20 mm longa, integra, sinuata, vel obtuse lobata. In-
florescentia paniculata, ramis flexuosis; Bores aggregati. Tepala 4, erecta,
naviculiformia, breviter pilosa; stamina 0-2, filamento non pulvinato. Pericarpium
diaphanum ad semen adherens, glabrum vel minute glandulosum. Semen erectum,
lenticulare, c. 0.6 mm altum; testa minute puncticulatum; embryo hippocrepiformis
ad apicem semine positus; radicula inferior.

Typus: 24 km E of Depot Springs HS., 28°54’S, 121° 20’E, Western Australia; base
of breakaway; 27 Aug. 1970, P. G. Wilson 8909 (holo: PERTH; iso: K).

Delicate erect aromatic annual herb to 30 cm high, moderately pilose all over with
weak gland-tipped and stiffer simple segmented hairs. Stem and branches slender.
Leaves alternate; lamina thin ovate to elliptic, 10-20 mm long, entire, sinuate, or ob-
tusely lobed; petiole slender c. half length of lamina. Inflorescence paniculate
eventually occupying most of plant, composed of open dichasia with flexuose
branches which become abbreviated towards apex of plant, the flowers sessile in
small clusters. Flowers bisexual and female, c. 1 mm high (c. 1.5 mm in fruit); tepals

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4, erect, naviculiform, shortly pilose, fleshy with membranous margins; stamens 0-2,
slightly exceeding perianth, filament linear (not pulvinate at base), anther broadly
cordate c. 0.25 mm long; stigmas 2 filamentous, united into a short style. Utricle en-
closed in perianth which remains herbaceous in fruit; pericarp dull, diaphanous and
adherent to seed, glabrous, or minutely glandular. Seed erect, lenticular, c. 0.6 mm
high; testa minutely puncticulate; embryo hippocrepiform around apex of seed, rad-
icle inferior. Fruit shed while enclosed in perianth.

Distribution. In the central portion of the Eremaean Botanical Province of Western
Australia. Map 15.

Habitat. On rocky outcrops and at the base of breakaways.

WESTERN AUSTRALIA: Mt Magnet, Sept. 1903, W. V. Fitzgerald (NSW); Pass of
the Abencerrages, Rawlinson Ra., A. S. George 12137 (PERTH); near Mt Squires, 27
Aug. 1891, R. Helms (AD).

Superficially this species would appear to be related to the 4-merous species in the
genus Dysphania (e.g. D. rhadinostachya) but in seed character it is obviously differ-
ent, being very similar to Chenopodium pumilio and its allies of the section
Orthosporum. Yet even here its placement is uncertain since the other species of this
section have flowers arranged in compact glomerules, have one stamen, and normally
a 5-merous perianth. Aellen (1939 in sched.) was of the opinion that it was a member
of the Chenopodium botrys complex, but those species (and others of the section
Botryoides) differ in having 5-merous perianths and horizontal seeds. In 1956
Brenan described the sect. Margaritaria which contained the single species Cheno-
podium congolanum (Haum.) Brenan; this species is similar to C. saxatile except
that it has whitish sub-globose hairs (as well as glandular) and only one stamen, in
addition the terminal flower of each axillary cyme has a horizontal (not vertical)
seed. Thus it appears that C. saxatile does not fit satisfactorily into any of the pres-
ently recognised sections. Rather than create a further section, which would be
monotypic, I have expanded the circumscription of the section Orthosporum to in-
clude this species.

The axillary inflorescence (a dichasium) is similar in morphology to the compact
glomerule typical of the sect. Orthosporum', its different appearance being due to the
more open lateral branches which, towards the base of the plant, themselves branch
and become flexuose.

The glandular hairs found in C. saxatile appear to be modifications of the normal
multicellular trichome; the gland itself being very small and often scarcely observ-
able. This situation is very different to that in the other species of the section where
the glandular hairs consist of a large globular gland cell, either sessile or on a short
stipe.

The specific epithet refers to rocky situations in which Chenopodium saxatile is
found.

2. DYSPHANIA R. Br.

R.Bj., Prod. 411 (1810); Moq. in DC., Prod. 13/2:86 (1849); Benth., FI. Austral. 5:164
(1870); J. D Hook in Bentham et J. D. Hook., Gen. PI. 3:14 (1880) “Illecebraceae”;
Payn Engler et Prantl, Nat. Pflanzenf. 3, lb:92 (1889) “Caryophyllaceae”; Pax et
Holtmann in Engler et Prantl, Nat. Pflanzenf. ed. 2, 16c:272 (1934)

Paul G. Wilson, Tribe Chenopodieae

181

“Dysphaniaceae”; Aellen in Hegi, III. FI. Mitt.-Eur. ed. 2, 3:748 (1961)
“Dysphaniaceae”; Eckhardt in Melchior, A. Engler’s Syllabus der Pflanzenf. ed. 12, _
2:96 (1964) “Dysphaniaceae”; Airy Shaw in Willis, Diet. FI. PL Ferns ed. 7.387 (1966)
“Dysphaniaceae”; A. J. Scott Bot. Jahrb. Syst. 100:217 (1978). — Chenopodium sect.
Dysphania (R.Br.) Aellen, Bot. Jahrb. Syst. 63:486 (1930); J. Black, Trans. Roy. Soc.
S. Austral. 58:173 (1934). Type.' D. littoralis R.Br.

Chenopodium sect. Tetrasepala Aellen, Bot. Jahrb. Syst. 63:490 (1930); J. Black,
Trans. Roy. Soc. S. Austral. 58:172 (1934). — Dysphania sect. Tetrasepalae (Aellen) A.
J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Lectotype: C. inflatum Aellen (see A. J.
Scott loc. cit.).

Dysphania sect. Caudatae A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Type; D.
plantaginella F. Muell.

Annual or short-lived aromatic perennial herbs with subsessile glands or simple
and gland-tipped multicellular hairs, aromatic. Leaves alternate, simple, entire or
variously dissected, fleshy or thin. Inflorescence of compact cymose glomerules of
flowers (thyrses), these either axillary, spicoid or paniculate in arrangement. Flowers
ebracteate, minute (0.5 mm long), bisexual and female; bisexual flowers (often
functionally male) terminal to the glomerule or to each side of the glomerule; female
flowers lateral. Tepals 1-4 (3 or 4 in terminal flowers of glomerule), free or united at
base; limb cucullate and often inflated of spongy. Stamens 1-2, anterior-lateral in
position, somewhat exceeding tepals at anthesis; filament slender thickened at base,
disc absent. Ovary ellipsoid, erect; stigmas 1-2, linear, delicate. Fruit a nut; pericarp
diaphanous, smooth to papillate. Seed erect, oblique, or horizontal, globular to ellip-
soid or laterally compressed; testa crustaceous, smooth; embryo lateral or basal, situ-
ated beneath a groove (“embryo-groove”), radicle either inferior or superior.

Ten species all endemic to mainland Australia.

The taxonomic position of Dysphania has been a matter of doubt or disagreement
amongst botanists since Brown (1810) first described the genus. Brown considered it
to have affinity to the family Chenopodiaceae, in which he placed the genus but evi-
dently with reservations. Moquin (1849) also included it in the Chenopodiaceae (as
Salsolaceae) and suggested affinities with Monolepis and Blitum. Bentham (1870)
suggested that it was closely related to the section Orthosporum of Chenopodium (a
view with which I concur). J. D. Hooker (1880) included it in the Illecebraceae and
Pax (1889), who was evidently influenced by J. D. Hooker, in the Caryophyllaceae.
However, Pax (1927) and later Pax and Hoffmann (1934) placed it in its own family,
the Dysphaniaceae, which they considered to be intermediate in morphology be-
tween (Chenopodiaceae and (Caryophyllaceae. This systematic arrangement was
adopted by Aellen (1961), Eckhardt (1964 loc.cit.) and Airy Shaw (1966), who all rec-
ognised its affinity to the Chenopodiaceae. Aellen had earlier (1930) made
Dysphania a section of Chenopodium as did Black (1934 l.c.).

The close relationship between Chenopodium and Dysphania appears to have
always been recognised by workers in Australia who were familiar with some endemic
species of the former genus that were very similar in habit and floral morphology to
those of the latter. This relationship was not so evident to European systematists
who would have compared species such as Chenopodium album with Dysphania and
found them to be very different. The confusion created by the rather extreme pos-
itions taken by several workers was finally resolved by Eckardt (1967, 1968, 1969)
and by Scott (1978b). Eckardt showed that the floral morphology of Dysphania cor-
responds to that of Chenopodium hut is quite different to that found in the

22493 — ( 4 )

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Nuytsia Vol. 4, No. 2 (1983)

Illecebraceae (or Caryophyllaceae). Even so the circumscription of Dysphania re-
mained unsatisfactory for it included only species with three or fewer perianth seg-
ments. The addition to the genus by Scott (1978b) of the two species placed by
Aellen (1930) in Chenopodium sect. Tetrasepala was a big improvement for it then
became a discrete taxon clearly separable from Chenopodium. Scott recognised three
sections in Dysphania; these were based on the number of perianth segments (1-2, 3,
or 4) and the arrangement of the flower clusters (axillary or spicate). These sections
appear to be natural ones (although the tepal numbers given by Scott are not always
correct) but with so few species it appears to be unnecessary to provide an
infrageneric classification.

The genus Dysphania is closely related to Chenopodium sect. Orthosporum (as
was recognised by Bentham, 1870). This section has similar glandular hairs and
glomerules of small flowers with 4-5 tepals and one stamen. The only characters
which suggest that the two genera should be recognised as distinct are found in the
seed. In Dysphania the seed (when erect) has either a basal or lateral embryo (or it is
intermediate in position); when lateral the radicle is either inferior or superior. In
Chenopodium sect. Orthosporum the erect, lenticular seed always has a hippocrepi-
form embryo curved around its apex. The position of the embryo in the Dysphania
seed, is indicated by a corresponding depression in the testa; this I am referring to as
the embryo-groove. The radicle is situated in a pouch formed by an invagination of
the testa; this pouch is usually slightly protruding and, since at this point the testa is
thicker, it also appears darker in colour. The orientation of the embry-groove and the
position of the pouch are characters useful in recognising the different species.

Key to species

1. Tepals 1-3

Tepals 4

2. Flowers in glomerules axillary to foliage leaves 3

Flowers in spike-like inflorescences 8

3.

4.

5.

6.

7.

8.

Tepals (of lateral flowers of glomerules) 1-2; seed erect, ellipsoid to obovoid or

laterally flattened

Tepals 3, seed napiform to subglobular 5. D. littoralis

Tepals united at base and shed with the enclosed seed 5

Tepals free; seed and tepals shed separately 8

Seed lenticular, embryo curved around its base 3. D. valida

Seed obvoid or ellipsoid, embryo lateral 1. D. glomulifera

Seed strongly flattened (usually slightly twisted), obovate to deltoid; pericarp

almost smooth. piatycarpa

Seed ellipsoid, obovoid, or lenticular; pericarp smooth or glandular-papillate . . 7

Pericarp smooth, puncticulate, or minutely papillate; seed ellipsoid to obovoid

or lenticular 1. D, glomulifera

Pericarp prominently papillate; seed asymmetrically narrow-obovoid (the em-
bryo side flat) 2. D. glandulosa

Seed compressed globular, embryo oblique to erect 7. D, sphaerosperma

Seed ellipsoid, erect q

Paul G. Wilson, Tribe Chenopodieae

183

9. Tepals free from each other, rounded or inconspicuously keeled on

back 6. D. plantaginella

Tepals united into a hard cup-shaped tube in lower half, prominently (and hori-
zontally) keeled 8. D. simulans

10. Plant prostrate or decumbent, many stems arising from base; seed (i.e. pericarp)

glossy, compressed globular; inflorescence slender spicate 10. D. kalpari

Plant erect with one main stem; seed dull (rarely glossy), almost globular; inflor-
escence paniculate 9. D. rhadinostachya

1. Dysphania glomulifera (Nees) Paul G. Wilson, comb. nov. — Atriplex
glomulifera Nees in Lehm., PI. Preiss 1:634 (1844); Moquin in DC., Prod. 13/2:103
(1849); Benth., FI. Austral. 5:177 (1870); Ulbrich in Engler et Prantl, Nat. Pflanzenf.
ed. 2, 16c:515 (1934). Type: “In cultis ad caput fluvii Cygnorum, Januario a 1840”.
Preiss 1257 (holo: LD; iso:P).

D. myriocephala Benth., FI. Austral. 5:165 (1870).— C. myriocephalum (Benth.)
Aellen, Bot. Jahrb. Syst. 63:488 (1930). Lectotype: Western Australia, J. Drummond
206 (holo: K n.v.; iso: MEL, P) lecto. nov.

D. benthamiana Domin., Biblio. Bot. 89:103 (1925). Type: “Flooded ground south of
Wills Creek, Howitts Expedition”, Dr Murray (iso: MEL 19878).

Prostrate, decumbent, or erect annual or short-lived perennial herb c. 25 cm diam.,
to 15 cm high, numerous slender stems arising from base, sparsely covered with sess-
ile or stipitate gland-tipped hairs. Leaves fleshy, crowded towards the apex; lamina
narrow to broad elliptic, (1)5(10) mm long, passing into a slender petiole c. half its
length. Glomerules of flowers dense, present in most leaf axils. Terminal flower (of
glomerule) bisexual; tepals 3, obovate, cucullate, c. 0.7 mm long, green, fleshy except
for scarious margin, glabrous or with a few marginal glandular hairs, shortly united
at base or free; stamens 2 shortly exserted at anthesis; style solitary or paired. Lat-
eral flowers (of glomerule) female; tepals 1 or 2 with a white bladdery strongly cucul-
late limb and a filamentous to linear claw; stamens absent; ovary ellipsoid, delicate;
styles 1 or 2. Fruiting perianth: tepals 0.7-1 mm long; limb inflated and fungoid
covering the fruit. Pericarp diaphanous dull, closely adhering to seed, puncticulate,
papillate, or granulate. Seed erect, bluntly ellipsoid, compressed obovoid, or some-
what lenticular with rounded margins, c. 0.5 mm long; testa smooth reddish brown;
embryo slightly curved extending along the dorsal lateral margin beneath the em-
bryo-groove; radicle inferior with the radicular pouch forming a small dark protuber-
ance at base of seed.

Typification. The type of Dysphania glomulifera (i.e. Preiss 1257) came from near
Guildford (c. 15 km east of Perth) in cultivated ground near the Swan River (‘in
cultis ad caput fluvii Cygnorum’). It is a precise match with the lectotype of D.
myriocephala (Drummond 206) which probably came from the same general area
since it is only near here that the particular variant this collection represents has
been found. Aellen (1938) recognised that the plant described by Nees (1844) as
Atriplex glomulifera did not belong to that genus but since Aellen assumed the name
to be a synonym of D. littoralis R.Br. (1810) he did not make a new combination
under Dysphania. Very few replicates of Preiss no. 1257 appear to exist; neither
Moquin (1849) nor Bentham (1870) saw this number (they relied on Nees’ descrip-
tion when writing their own works) and both assumed the species to be correctly
placed in Atriplex although a puzzling member. Moquin (1849) based his descrip-
tion of D. littoralis on Drummond 206, the lectotype of D. myriocephala; therefore
his description applies to D. glomulifera rather than to the former species.

184

Nuytsia Vol. 4, No. 2 (1983)

Key to subspecies

Seed compressed obovoid to bluntly lenticular; pericarp smooth (rarely

papillate)

Seed ellipsoid; pericarp granulate

la. subsp. glomulifera
lb. subsp. eremaea

la. subsp. glomulifera Figure 6A.

D. myriocephala Benth., op. cit . — Chenopodium myriocephalum (Benth.) Aellen, op.
cit.

Lateral flowers of glomerules: tepals 1(2); claws filiform to linear, readily
separating from pedicel (and from opposite member where present); styles (1)2. Seed
broadly compressed obovoid to bluntly lenticular; pericarp smooth or minutely
puncticulate (rarely papillate).

QUEENSLAND: Moonie Ponds, S. L. Everist 3718 (BRI); Cunnamulla, P. Martensz
62 (CANB).

NEW SOUTH WALES: Near Booligal, E. M. Canning 3859 (CBG); Coolabab, W. E.
Mulham W857 (NSW).

VICTORIA: 15 mi [24.1 km] NE of Hattah, A. C. Beauglehole 40569 (PERTH)- Lake
Eildon, 14 May 1973, J. H. Willis (MEL).

SOUTH AUSTRALIA: Loxton, Hj. Eichler 12326 (AD); Lenswood, A. G. Spooner

1 *7 A / A \ ■*

5174 (AD).

WESTERN AUSTRALIA: Guildford, 12 Aug. 1901, C. R. P. Andrews (NSW)-
Leederville, Mar. 1967, 1. S. Hutchinson (PERTH).

Distribution. Near Perth in Western Australia, south-eastern and north-eastern
South Australia, Victoria, eastern New South Wales and Queensland. Map 17.

Habitat. Principally along banks of freshwater streams or rivers and in muddy areas.
Some ecotypes in forest areas, rocky hillsides, or as weeds of cultivation.

Ib. subsp. eremaea Paul G. Wilson, subsp. nov. Figure 6B.

D. benthamiana Domin., op. cit.

Flores laterales: tepala 2, limbis multo inflatis fructum obtegenti; stylus
plerumQu© solitarius. Semen obtuse ellipsoideumj pericsrpium granulatum.

Birdsville, S. T. Blake 12224 (BRI); Boorara Stn., R. C. Carolin

4035 (SYD).

Paul G. Wilson, Tribe Chenopodieae

185

NEW SOUTH WALES: Wansaring, L. A. S. Johnson 547/125 (NSW); Wanheroo
Hills, P. G. Milthorpe 1915 (NSW).

SOUTH AUSTRALIA: Coglin River, Basendow 7 (AD); Millars Ck. Stn., R. J.
Chinnock 2573 (AD).

WESTERN AUSTRALIA: 22 mi [35.4 km] W of Browne Ra., A. S. George 5417
(PERTH); 18 mi [29 km] E of Meekatharra, T. E. H. Aphn 2460 (PERTH).

NORTHERN TERRITORY: Redbank Gorge, G. Chippendale (N T 4878); Mt
Doreen Stn., P. K. Latz 2002 (NSW).

Distribution. South-West and central Queensland, north-west New South Wales,
northern South Australia, central Western Australia, southern portion of the North-
ern Territory. Map 17.

Habitat. Damp sand near freshwater.

The typical variants of the two subspecies of D. glomulifera are quite distinct (at
least as to microscopic characters) but there are numerous other variants which blur
the separateness and provide additional characters that extend the morphological
complexity. Since the distinguishing features are so small (and fragile) and appear to
be subject to environmental modification, it appears unwise to treat the two taxa at
the species level or to formally recognise further infraspecific groups.

Near the Murray River in Victoria are found two variants of subsp. glomulifera-, in
one the pericarp is smooth and in the other it is papillate. There do not appear to be
other morphological characters which might support the recognition of these vari-
ants as distinct taxa. The variant with papillae on the pericarp may be readily dis-
tinguished from D. glandulosa by the shape of the seed.

The variant of subsp. glomulifera found in central and eastern New South Wales
and in central and eastern Queensland, has a bluntly lenticular seed. This variant
may be clearly separated from D. valida by its free (or almost free) tepals and by the
embryo being lateral, not basal, in position.

The subspecies eremaea represents the plant referred by Aellen (1930) to Cheno-
podium blackianum, and by J. M. Black (1948) to its nomenclatural synonym
Dysphania littoralis. Most Australian herbaria have followed Black in their misappli-
cation of the latter name. Domin (1925) appears to have recognised the two taxa as
distinct and described as a new species, Dysphania henthamiana, the plant here
called subsp. eremaea. Unfortunately Domin incorrectly made D. myriocephala a
synonym of D. littoralis (as ‘D. litoralis’) although he cited no material of that taxon;
he also failed to recognise the critical characters which distinguished the species in
this group. Dysphania glomulifera subsp. eremaea is, in fact, quite distinct from D.
littoralis for in the latter species all flowers have three tepals firmly united at their
base and deciduous together with the enclosed fruit, while the seed is sub-globular
and the embryo horizontal.

The illustration of ‘Dysphania litoralis’ provided by F. Mueller (1889-1891 t. 36),
was evidently intended to portray the plant here referred to D. glomulifera subsp.
eremaea. Unfortunately it would appear that the artist either exercised considerable
licence or obtained material from several species and combined their different fea-
tures in the one plant. In addition, the seed has been drawn incorrectly; the embryo

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is indicated as being on the opposite side to that of the longitudinal groove (whereas
it lies beneath it) and the radicular pouch is placed in an apical position (instead of
basal) although the radicle itself is correctly illustrated as being inferior.

The infraspecific epithet is derived from the Greek word eremia, a desert, alluding
to the typical habitat of this plant.

2. Dysphania glandulosa Paul G. Wilson, sp. nov. Figure 6C.

Herba prostrata; caules numerosi graciles ad basim exorientes. Folia parva, lamina
anguste elliptica c. 3 mm longa. Flores in glomeratis axillaribus in fructu c. 1 mm
diam. Flos terminalis hermaphroditus; tepala 3, libra; stamina 2; styli 2. Flores lat-
erales feminei; tepala (1)2, libra, c. 0.5 mm longa, delicatissima, fructu imperfecte
obtegentia. Pericarpium diaphanum, papillis prominentibus glanduliformibus
ornato. Semen asymmetrice semi-obovoideum (latere embryonis piano), c. 0.3 mm
longum, apice rotundato.

Typus: 28 km NE of Nambi Homestead which is 65 km NNE of Leonora (28°8'S,
121°43'E); prostrate in red loam; 29 Aug. 1968, P. G. Wilson 7503 (holo: PERTH).

Prostrate annual herb c. 20 cm diam., numerous slender stems arising from base
and giving rise to many lateral branches, sparsely pilosulose all over (except inflor-
escence) with simple and glandular multicellular hairs. Leaves small, slightly fleshy;
lamina narrow-elliptic, c. 3 mm long; petiole equal to lamina. Inflorescence of
axillary glomerules as in D. glomulifera; glomerules (in fruit) c. 1 mm diam. Ter-
minal flower of glomerule bisexual; tepals 3, free, the two anterior-lateral 0.4 mm
long in fruit and larger than the posterior; stamens 2 opposite anterior-lateral tepals;
styles 2, slender, c. 0.1 mm long. Secondary terminal flowers (when present) similar
to terminal but with two tepals and one stamen. Lateral flowers female, tepals (1) 2,
free, 0.5 mm long (in fruit), very delicate and incompletely covering fruit; limb
strongly cucullate, slightly inflated, emarginate with a scarious margin; claw linear
± equal to limb. Pericarp diaphanous, covered with prominent gland-like papillae.
Seed erect, asymmetrically semi-obovoid (the embryo-side flat), c. 0.3 mm long,
apex rounded; testa smooth, reddish brown; embryo-groove longitudinal along flat-
tened side of seed; embryo lateral, erect, radicle inferior. Infructescence
disintegrating at maturity into separate tepals and fruits.

WESTERN AUSTRALIA: Glenorn Street, August 1938, N. T. Burbidge (PERTH);
Belele Station, D. W. Goodall 3351 (PERTH).

Distribution. West-central area of Western Australia. Map 18.

Habitat. Loam; seasonally waterlogged areas.

Dysphania glandulosa may be distinguished from D. glomulifera (to which it ap-
pears to be most closely related) by its relatively small leaves, its small tepals (which
do not completely cover the apex of the fruit), its strongly papillate pericarp, and by
the shape of its seeds. One variant of D. glomulifera subsp. glomulifera also has pa-
pillae on the pericarp but in this plant the seed is broadly compressed-obovoid or len-
ticular whereas in D. glandulosa it is asymmetrically obovoid (flattened on the side
bearing the embryo-groove).

The specific epithet alludes to the gland-like appearance of the papillae on the
surface of the pericarp.

Paul G. Wilson, Tribe Chenopodieae

3. Dysphania valida Paul G. Wilson, sp. nov.

187

Herba perennis, prostrata, caulibus aliquot per radicem orientibus. Folia
crassiuscula; lamina anguste oblonga vel anguste elliptica, integra, 10-15 mm longa.
Flores in glomeratis axillaribus 3-5 mm diam. Flos terminalis masculinus; tepala 3 ad
basim breviter connata; stamina 2; pistillodium minutum. Flores laterales feminei;
tepala 2, in fructu c. 1.3 mm longa, ad basim breviter connata, limbo inflato ob-
ovoideo cucullato, unguide breviter oblongo; styli 2. Pericarpium diaphanum laeve.
Semen erectum, lenticulare, c. 0.6 mm altum; embryo semicircularis basalis; radicula
cotyledonesque erectae.

Typus: Warrego District, near Morven; hard red box-sandalwood country; common;
6 April 1941; C. T. White 11895 (holo: BRI 061936).

Short-lived herbaceous perennial, prostrate, to 60 cm diameter, almost glabrous to
sparsely pilose with short, multicellular, simple or gland-tipped hairs. Stems several,
slender, arising from rootstock. Leaves somewhat fleshy; lamina entire, narrow-ob-
long to narrow-elliptic, obtuse, 10-15 mm long; petiole slender c. half length of
lamina. Glomerules of flowers dense, axillary, 3-5 mm diam. Terminal flower (of
glomerule) functionally male; tepals 3 shortly united at base; stamens 2 opposite an-
terior-lateral tepals; pistillode minute, c. 0.2 mm high. Lateral flowers (of glomerule)
female; tepals 2, at fruiting stage with an inflated obovoid cucullate limb scarious at
the margin and a short broad claw united to opposite member at base, in all c.
1.3 mm long, glabrous or sparsely pilose with gland-tipped hairs; styles 2, short and
delicate. Pericarp diaphanous, smooth, adherent to seed. Seed erect, lenticular, c.
0.6 mm long; testa smooth; embryo semicircular occupying lower 2/3 of seed; radicle
and cotyledons erect; raphe extending from apex to base of seed. Infructescence
breaking up at maturity to release fruit enclosed in the persistent tepals.

QUEENSLAND: Morven, S. T. Blake 10914 (BRI); Eulo, August 1967, M. Law
(BRI).

Distribution. Southern Queensland (south of 26° latitude). Map 19.

Habitat. Collections have been noted as coming from “cultivation paddock in forest
soil”, hard red box -sandalwood country, and “timbered, rocky, sandy hilltop”.

This species is similar to D. littoralis in its habit and in its firmly united tepals; it
differs most obviously in the female flowers possessing only two tepals, in the len-
ticular seed and in the basal embryo. From D. glomulifera it differs in having larger
tepals and again in its seed shape and embryo position.

The specific epithet valida is a Latin word meaning strong, and alludes to the
firmly united tepals.

4. Dysphania platycarpa Paul G. Wilson, sp. nov. Figure 6D.

Herba prostrata sparse pilosulosa. Folio fere glabra; lamina elliptica 5-10 mm
longa. Flores in glomeratis axillaribus in fructu c. 2-5 mm diam. Flos terminalis
hermaphroditus; tepala 3, libra, c. 1 mm longa, prominente cucullata; stamina 2; styli
2, perbreves. Flores laterales feminei; tepala 1 vel 2, c. 0.7 mm longa. Semen erectum,
applanatum, obovatum vel deltoideum, c. 0.4 mm longum, leviter tortum vel
lateribus concavis; embryo lateralis, radicula inferior!. Pericarpium minute
granulatum.

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Nuytsia Vol. 4, No. 2 (1983)

Typus: West of Western Australia — Northern Territory border at 23°10'S; edge of
claypan, prostrate, 26 July 1967, A. S. George 8927 (holo: PERTH).

Prostrate annual herb to 20 cm diam., sparsely pilosulose with simple or glandular
multicellular hairs, numerous slender mostly simple stems arising from base. Leaves
slightly fleshy, almost glabrous; lamina elliptic, obtuse, 5-10 mm long; petiole slen-
der, ± equal to lamina. Glomerules of flowers dense, axillary, c. 2.5 mm diam. in
fruit. Terminal flower of glomerule bisexual; tepals 3, free, c. 1 mm long, prominently
cucullate, claw filiform; stamens 2 opposite and ± equal to anterior-lateral tepals;
styles 2, very short; secondary terminal flower similar to but slightly smaller than
terminal with two tepals and one stamen. Lateral flowers female; tepals 1 or 2,
strongly cucullate in fruit with delicate white inflated limh and a filamentous claw,
in all c. 0.7 mm long. Pericarp diaphanous, minutely granulate, intimately adherent
to seed. Seed erect, obovate to deltoid, flat, c. 0.4 mm long, slightly twisted or with
faces sunken; testa dull reddish brown; embryo lateral, radicle inferior.
Infructescence readily breaking up when mature into separate tepals and seeds.

QUEENSLAND: Birdsville, S. T. Blake 12224 (BRI); Currawilla, S. L. Everist 3977
(CANB).

NEW SOUTH WALES: Broken Hill district, A. Morris 890 (ADW, NSW); Sturt
Nat. Park, W. E. Mulham W937 (NSW).

SOUTH AUSTRALIA: 13 km NW of Quinyambie H.S., N. N. Donner 3593 (AD);
Abminga, E. H. Ising 2397 (AD).

WESTERN AUSTRALIA: W of W.A.— N.T. border at 23°10’S, A. S. George 8927
(PERTH).

NORTHERN TERRITORY: Mulga Park Stn., T. S. Henshall 134 (MEL); Andado
Stn., PK Late 6826 (NT).

Distribution. Central Australia; South-west Queensland, extreme western New South
Wales, central, northern and north-east South Australia, extreme west-central West-
ern Australia, south eastern Northern Territory. Map 20.

Habitat. In clay or mud by fresh water.

Dysphania platycarpa may be readily distinguished from related species (i.e. D.
glomulifera, D. glandulosa, and D. littoralis) by the slender claws to the tepals and
by the peculiar fruit. This fruit, unlike that of all other Dysphania species, is flat but
slightly twisted (or sunken on the hroad faces), and the pericarp is minutely granu-
late.

The specific epithet is derived from two Greek words platy, broad, and carpos,
fruit, with reference to the shape of the seed.

5. Dysphania littoralis R.Br., Prod. 412 (1810). -Chenopodium blackianum
Aelien, Bot. Jahrb. Syst. 63:487 (1930) based on above, non C. littorale (L.) Thunb.
(1815). Type: Port I [near Gladstone], Queensland, 6 August 1802, R. Brown (holo:
BM; iso: K), see comment below. Figure 7B.

Prostrate annual (or perennial?) herb with fleshy taproot, c. 30 cm diam., sparsely
pilosulose with simple or gland-tipped multicellular hairs, numerous slender, mostly
simple, radiating stems arising from base. Leaves somewhat fleshy; lamina elliptic

Paul G. Wilson, Tribe Chenopodieae

189

obtuse, 5-10 cm long, petiole slender somewhat shorter than lamina. Glomerules of
flowers axillary, dense, c. 2 mm diam. Terminal flower (of glomerule) male; tepals 3,
one posterior and two anterior-lateral, shortly united at base, obovate, the limb in-
flated, in all c. 1.3 mm long; stamens 2 opposite the anterior-lateral tepals, anther c.
0.3 mm long; pistillode minute c. 0.2 mm high with a pair of short styles. Secondary
terminal flower (when present) similar to the terminal but with only one stamen.
Lateral flowers female; tepals 3 (4), at fruiting stage obovoid, very inflated (fungoid),
without a scarious margin, white, claw narrow oblong somewhat shorter than the
limb, in all c. 0.8 mm long; style 1 (rarely 2), slender. Pericarp diaphanous, smooth,
glossy, closely adherent to the testa. Seed napiform to depressed globular, horizontal
to oblique, 0.3-0. 4 mm high, slightly furrowed over the horizontal embryo; testa
smooth, reddish brown, raphe extending from radicle to base of seed. Infructescence
breaking up when mature to release the seed enclosed in the three persistent tepals.

QUEENSLAND: Gladstone, A. Dietrich 2417 (MEL); Curtis Is., S. T. Blake 22525
(BRI).

NEW SOUTH WALES: Idalia, South Wanaaring, P. L. Milthorpe and G. M. Cun-
ningham 4768 (NSW).

WESTERN AUSTRALIA: Upper Murchison, 1890, /. Tyson (MELU).

Distribution. Northern New South Wales; south eastern and east coastal
Queensland; possibly the Murchison River district of Western Australia (see com-
ment below). Map 18.

Habitat. The coastal variant in tidal mud flats; the inland variant in loam in open
forest.

The name Dysphania littoralis has been used loosely to include D. glomulifera or,
when the latter species has been recognised (usually as D. myriocephala), has been
applied to the inland plant here referred to D. glomulifera subsp. eremaea. The two
species are quite distinct: in D. littoralis the lateral flowers have three (rarely four)
tepals which remain united at their base and the seed is turnip-shaped with a hori-
zontal to oblique embryo; in D. glomulifera (and in the related species D. glandulosa,
D. platycarpa, and D. valida) the lateral flowers have one or two tepals and the seed
and embryo are erect.

The illustration under the name ‘D. litoralis’ provided by F. Mueller in his Icon.
Austral. Salsol. PL tab. 36 (1890) was evidently derived from several collections,
possibly none of which were of this species. For further discussion see under D.
glomulifera..

The specimen collected by Isaac Tyson from the upper Murchison River is the
only record of this species from Western Australia, while it has not been recorded
from the Northern Territory, South Australia, or western Queensland.

The type of Dysphania littoralis was collected by Robert Brown on coastal mud-
flats in the tropics near Gladstone in Queensland. Material which matches the type
comes from such habitats in Queensland along its east coast. A variant very similar
to the type form has been collected at inland localities in southern Queensland and
north-west New South Wales and here grows in quite different habitats such as
‘mixed open forest, sandy soil’ and ‘hard red soil’. It is possible that the inland

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Nuytsia Vol. 4, No. 2 (1983)

ecotype should be recognised as a distinct taxon but, since I am unable to find clear
morphological differences on which such a recognition could be based, I have in-
cluded both variants under the same name.

Typification. George Bentham (1870) stated that Robert Brown had collected the
type while running from some Aborigines, a comment that may relate to a note in
Brown’s unpublished diary in which he records an attack having taken place at
Keppel Bay on 16 August 1802. However, in his manuscript Flora, Brown states that
the collection was made at 'Port F on 6 August 1802, which information is also given
on the labels of the holotype and isotypes in the herbaria BM and K. Port I is near
the present port of Gladstone and about 50 km SE of Keppel Bay.

6. Dysphania plantaginella, F. Muell., Fragm. 1:61 {1858}.— Chenopodium
plantaginellum (F. Muell.) Aellen, Bot. Jahrb. Syst. 63:487 (1930). Lectotype: Sturts
Creek, Australia sub-centralis, March 1856, F. Mueller (MEL 19916) lecto. nov Fig-
ure 7E.

Annual herb with several prostrate to ascending stems arising from base. Stems
sparingly branched, pilose with multicellular hairs. Leaves sparsely pilose and with
sessile glandular hairs; lamina elliptic to broadly elliptic, entire, obtuse, 1-2 cm long,
base cuneate passing into a short petiole. Inflorescence of terminal, shortly peduncu-
late, very narrowly cylindrical usually erect spikes composed of continuous (or
interrupted) globular dichasial cymes; bracts solitary beneath each globular cluster,
with an inflated cucullate rostrate limb and a slender claw, in all c. 1 mm long.
Flowers with a slender pedicel 0.3-0.5 mm long. Terminal flower (of dichasial cyme)
bisexual; tepals 3, free but remaining attached to pedicel, erect, limb concave and
strongly inflated with a rounded to spreading keel, ciliolate otherwise glabrous,
closely enveloping seed; stamen solitary opposite an anterior-lateral tepal; style slen-
der, solitary. Lateral flowers (of dichasial cyme) numerous, female, otherwise similar
to terminal flower. Pericarp diaphanous, faintly areolate otherwise smooth. Seed el-
lipsoidal, erect, c. 0.5 mm high; embryo erect, lateral and basal beneath a longitudi-
nal groove, the radicle inferior; testa smooth, reddish brown. Infructescence breaking
up when mature at base of pedicels to release seed enclosed within the three tepals.

NEW SOUTH WALES: 23 km NNW of Conoble Rly Stn., J. Pickard 1943 (NSW).

SOUTH AUSTRALIA: Koonamore Stn., Hj. Eichler 17177B (AD); 2 mi [3.2 km]
SE of Oodnadatta, A. C. Beauglehole 20839 (AD).

WESTERN AUSTRALIA: Nita Downs Stn., R. A. Saffrey 1680 (PERTH); 32 km W
of Gascoyne Junction, P. G. Wilson 8424 (PERTH).

NORTHERN TERRITORY: 30 mi [48.3 km] N of Willowra HS, G. Chippendale
(NT4739); 6 mi [9.7 km] S of Hermannsburg Mission, P. K. Latz 793 (AD).

Distribution. Central New South Wales north westwards through central and north-
ern South Australia, the Northern Territory to northern and north western Western
Australia. Map 18.

Habitat. Sand, either coastal or on the margin of waterholes and inland saltlakes.

Dysphania plantaginella is rather variable in its morphology, the dichasial cymes
sometimes forming discrete globular clusters with rounded dorsal margins to the
tepals, and sometimes forming an apparently continuous spike, with slightly spread-

Paul G. Wilson, Tribe Chenopodieae

191

ing keels to the tepals. In addition, whereas most collections appear to have erect
spikes, in a few these are quite prostrate. These variants are not clearly defined so it
seems unwise to establish infraspecific taxa to accommodate them. In view of the
variety of habitats in which this species is found several ecotypes might be involved.

Dyspbania plantaginella is evidently closely related to D. sphaerosperma but dif-
fers most obviously in seed shape and in the nature of the pericarp. In D.
plantaginella the seed is ellipsoidal and the pericarp areolate (marked out in a fine
reticulate pattern) whereas in D. sphaerosperma the seed is compressed-globular
and the pericarp minutely granular-papillate.

7. Dysphania sphaerosperma Paul G. Wilson, sp. nov. Figure 6E-F.

Herba prostrata vel ascendens; caules numerosi ad basim exorientes. Folia
pilosulosa; lamina elliptica obtusa integra vel undulata 0.5-1. 5 cm longa.
Inflorescentia spicata; spicae breviter pedunculata, anguste cylindraceae ad 15 cm
longae, cymarum congestorum compositae. Flores fere sessiles vel pedicello crasso c.
0.1 mm longo instruct!. Flos terminalis hermaphroditus (vel femineus); tepala 3, c.
1mm longa, libra; stamen solitarium; stylus solitarius gracilis. Pericarpium
diaphanum, minute granulatum. Semen compresse globulare c. 0.5 mm alte; embryo
semicircularis obliques vel erectus, radicula inferiore. Tepala fructum cingentia per-
sistentia.

Typus: Upper reaches of Turner River, Woodstock Stn., south of Port Hedland,
Western Australia; travertine ridge near river, ascendent herb, 28 April 1958, N. T.
Burbidge 5924 (holo: CANB 53651; iso: AD).

Annual herb with several prostrate to ascending stems arising from base. Stems
sparingly branched, pilosulose with multicellular hairs; lamina elliptic, obtuse, entire
or undulate 0.5-1. 5 cm long, passing into a short petiole. Inflorescence of terminal,
shortly pedunculate, narrowly cylindrical spikes to 15 cm long, composed of con-
gested dichasial cymes of 3 to several flowers; bracts (beneath each cyme) with an in-
flated cucullate rostrate limb and a slender claw, in all c. 1 mm long. Flowers almost
sessile or with a short thick pedicel c. 0.1 mm long. Terminal flower (of cyme) bisex-
ual (or female); tepals 3, c. 1 mm high in fruit, free but remaining attached to pedi-
cel, deeply cucullate and inflated with a rounded inflated limb and a short broad
claw, glabrous except for a few sessile glands towards the margins; stamen solitary
opposite an anterior-lateral tepal; style solitary, slender. Lateral flowers (of cyme)
similar to terminal flower but female. Pericarp diaphanous, minutely granulate pap-
illose. Seed compressed globular c. 0.5 mm high; embryo semicircular, oblique or
erect, lying beneath a deep median groove, the radicle inferior; testa smooth, reddish
brown. Infructescence breaking up when mature with flowers dehiscing at base of
pedicels releasing the seeds enclosed in the three tepals.

WESTERN AUSTRALIA: Wolf Meteorite Crater, A. C. Beauglehole 47376
(PERTH); Dirk Hartog Is., A. S. George 11413 (PERTH).

NORTHERN TERRITORY: Curtin Springs Stn., T. S. Henshall 728 (NT); Lake
Neale, P. K. Latz 2673 (NT).

Distribution. Shark Bay area on the west coast of Western Australia eastwards to the
southern area of the Northern Territory. Map 21.

Habitat. Possibly always associated with gypseous or calcareous soil, frequently
around salt lakes.

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Nuytsia Vol. 4, No. 2 (1983)

Dysphania sphaerosperina is similar in appearance to D. plantaginella-, it differs in
having rounded inflated tepals (not keeled), a very short thick pedicel, and
subglobular seeds with a minutely granulate-papillose pericarp. Several herbarium
collections consist of plants with only female flowers.

The specific epithet is derived from the Greek words sphaera, a sphere, and
sperma, a seed, referring to the almost globular seeds.

8. Dysphania simulans F. Muell. et Tate ex Tate, Trans. Roy. Soc. S. Austral.
8:71 (1886). — Chenopodium simulans (Tate) F. Muell., Sec. Syst. Census 50 (1889);
J. Black, Trans. Roy. Soc. S. Austral. 58:174 (1934). Type: Gypsum and salt flats in
the spring country, Cootanoorina, M. Murray (holo: AD; iso: MEL, NSW, PERTH).

C. osbornianum Aellen, Bot. Jahrb. Syst. 63:488 (1930). Type: “Koonamore Head
Station, 1914”, T. G. B. Osborn (holo: G n.v.; iso: AD96630144).

Annual herb with several decumbent to ascending branches arising from its base
and from below the inflorescences, 10-30 cm high, pubescent with simple
multicellular hairs on branches, petioles, and inflorescence axes, and sessile
glandular hairs on leaves and flowers. Leaves early deciduous; lamina chartaceous,
elliptic, 1-2 cm long, with 2-4 pairs of prominent obtuse lobes, apex obtuse to acumi-
nate, base cuneate and passing into a slender petiole somewhat shorter than lamina.
Inflorescence an erect narrowly cylindrical spike 5-20(30) cm long, 4 mm diam., sess-
ile at points of branching of stem, composed of densely arranged normally 7-flowered
dichasial cymes; bracts (beneath each cyme) with an inflated strongly cucullate ros-
trate limb and a slender claw, in all c. 1 ram long. Terminal flower (of cyme) large
and sessile, bisexual; tepals 3, cucullate, c. 1 mm long, united in lower half into an in-
durated strongly papillose cup-shaped tube; stamen solitary in axil of an anterior-
lateral tepal, sty'Ies 2, slender. Lateral flowers (of cyme) female otherwise similar to,
although smaller than, terminal flower. Fruiting perianth cartilaginous (especially at
the cup-shaped base), at first firmly attached to axis; tepal limbs produced into
spreading horizontal hyaline wings, in all 2 mm high and wide. Seed erect, broadly
pear-shaped, c. 0.6 mm high; embryo vertical extending along one side and around
base of seed; Radicle superior occupying the small dark-coloured terminal apiculum;
raphe extending from apex to base. Infructescence tardily breaking up, the seed en-
closed in the cartilaginous perianth which eventually splits into its separate
components.

QUEENSLAND: Lake Crocker, 28 June 1939, R. L. Crocker (AD); Poeppel Corner

D. E. Boyfand 237 (NSW).

NEW SOUTH WALES: 63 mi [101.4 km] SE of Wanaaring, W. E. Mulham W445

VICTORIA: 28.8 km W of Nowingi, 29 July 1979, J. M. Browne (MEL).

SOUTH AUSTRALIA: Koonamore Stn., M.D. Crisp 709 (AD); Oodnadatta, 8 Apr.
1950. E. E. Lord (MEL).

WESTERN AUSTRALIA: 37 km N of Agnew, D. Symon 9941 (ADW); 30 km NE of
Nambi HS., P. G. Wilson 7499 (PERTH).

TERRITORY: 11 km SW of Erldunda Stn., T. S. Henshall 46 (NT);
Mt Wedge Stn., P. K. Latz 2146 (AD).

Paul G. Wilson, Tribe Chenopodieae

193

Distribution. Central Australia; south-west Queensland, north-west New South
Wales, north-east South Australia, east central Western Australia, southern portion
of the Northern Territory. Map 21.

Habitat. Saline (halitic), gypseous, or calcareous soil, often on margin of salt lakes.

The densely arranged cymes form long terminal ‘mouse-tail’ inflorescences which,
because of their erect and sessile habit, give the plant a candelabrum aspect. This,
and the indurated cup-shaped base to the perianths, make Dysphania simulans a
very distinctive species.

The illustration by Graff provided by Mueller in his Icon. Austral. Salsol. PL tab.
34 (1890) well portrays the species; however, it shows the embryo incorrectly with
the cotyledons occupying the terminal apiculum whereas, in fact, this area is filled by
the radicle. The orientation of the embryo is thus the reverse to that found in D.
plantaginella (which has a similarly shaped seed); this is because the embryo differs
in its position by 180° to that which it occupies in D. plantaginella causing the hilum
to be moved near the apex of the seed and the raphe to elongate. The illustration
(tab. 34) showing an incorrect orientation of the embryo appears to have been di-
rected by Mueller from a specimen of A. Henry from Georgina River (MEL 19933)
for there are some pencil sketches by Mueller in which he evidently instructed Graff
on the fine detail of the fruit.

9. Dysphania rhadinostachya (F. Muell.) A. J. Scott, Bot. Jahrb. Syst. 100:218
(1978 ). — Chenopodium rhadinostachyum F. Muell., S. Sci. Rec. 2:98 (1882); F.
Muell., Icon. Austral. Salsol. PI. t. 33 (1890) p.p. Type: Finke River, 1881, H. Kempe
413 (holo: MEL 20098). Figure 7C-D.

Chenopodium rhadinostachyum var. paniculatum F. Muell. et Tate, Trans. Roy. Soc.
S. Austral. 16:345 (1896) nomen.

Chenopodium inflatum Aellen, Bot. Jahrb. Syst. 63:490 (1930). — D. inflata (Aellen)
A. J. Scott, Bot. Jahrb. Syst. 100:218 (1978). Lectotype: Bulloo Range, 29 August
1923, MacGillivray (holo: G n.v.; iso: ADW 17774, BRI 010647) lecto. nov.

Erect herb to 30 cm high with a single main stem, copiously branched above the
base, moderately pilose all over with short glandular and longer segmented hairs,
aromatic. Leaves thin; lamina elliptic, 10-20 mm long, undulate or with a few obtuse
lobes, base cuneate and passing into a slender petiole. Inflorescence of axillary and
terminal leafy or spike-like thyrses which occupy nearly the whole plant and are
composed of compact dichasial cymes of 3-7 flowers subtended by a bract. Bracts of
the inflorescence either foliaceous and shortly exceeding cymes or very reduced, nar-
rowly ovate acuminate c. 2 mm long. Pedicels very short, turbinate, c. 0.2 mm long,
all apparently arising from main axis. Flowers bisexual and female (the terminal
flower of cyme always bisexual, the lateral sometimes so); tepals 4, erect, c. 1 mm
long, incurved and cucullate, united in lower third, rounded or shortly and bluntly
horizontally keeled on back, inflated in fruit, glabrous to hirsute, ciliate; stamen 0-1,
about twice length of perianth; styles 2, slender, long exserted. Pericarp diaphanous,
dull, sometimes slightly scurfy, somewhat loosely adherent to seed. Seed subglobular,
horizontal to erect, c. 0.5 mm diam., slightly keeled and grooved over the semicircu-
lar embryo, radicle and cotyledons superior; testa reddish brown; raphe extending
from apex to base of seed. Infructescence breaking up with flower dehiscing at apex
of pedicel, the fruit loosely enclosed in perianth.

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Key to subspecies

Tepals hirsute 9a. subsp. rhadinostachya

Tepals glabrous or sparsely hirtellous, ciliate 9b. subsp. inflata

9a. subsp. rhadinostachya

Tepals hirsute, rounded on back or slightly keeled. Seed oblique to erect; pericarp
dull. Inflorescence much-branched with the bracts smaller than or exceeding the
glomerules.

QUEENSLAND: Woodstock, S. T. Blake 6499 (BRI); Mt Isa, R. L. Specht and R.
W. Rogers 30 (BRI).

SOUTH AUSTRALIA: Mt Ilbillee area, A. C. Beauglehole 25520 (AD); Mt Morris,
Hj. Eichler 17333(AD).

WESTERN AUSTRALIA: Dampier, 30 March 1971, H. Hukin (PERTH);
Rawlinson Range, P. Wilson 2378 (AD).

NORTHERN TERRITORY: Mt Olga, G. Chippendale (NT 4665); Mann Range, D.
J. Nelson 362 (NT).

Distribution. Western Queensland, northern South Australia, north-west to east-
central Western Australia, and southern half of the Northern Territory. Map 19.

Habitat. Typically a species of rocky slopes in skeletal soil.

9h. subsp. inflata (Aellen) Paul G. Wilson, stat. nov. — Chenopodium infiatum
Aellen, Bot. Jahrb. Syst. 63:490 (1930). Type: loc. cit.

Tepals glabrous or sparsely hirtellous, ciliate, rounded on back. Seed horizontal or
slightly oblique; pericarp dull (or rarely glossy). Inflorescence of long slender spikes
often with slightly disjunct glomerules; bracts inconspicuous.

QUEENSLAND: Boulia, S. T. Blake 6475 (BRI); 51 mi [82.1 km] W of Yaraka, L.
Everist 7368 (BRI).

NEW SOUTH WALES: Byrock, E. McBarron 14706 (NSW).

WESTERN AUSTRALIA: Mt Anderson Stn., R. D. Royce 6919 (PERTH); 74 km E
of Carnarvon, P. G. Wilson 8393 (PERTH).

NORTHERN TERRITORY: Stirling Swamp, T. S. Henshall 510 (NT); 110 km N of
Alice Springs. J. Z. Weber 995 (AD).

Distribution. Central Queensland, northern New South Wales, north-western West-
ern Australia, and southern half of the Northern Territory. Map 20.

Habitat. Evidently usually in skeletal soil but also recorded from ‘hard red mulga
country’ in Queensland.

Paul G. Wilson, Tribe Chenopodieae

195

Dysphania rhadinostachya has been confused with D. kalpari which may be dis-
tinguished by its decumbent habit and erect glossy seeds. The subspecies inflata
varies considerably in habit; part of this variation is due to age and environmental
conditions and part to the presence of several distinct forms which are not recog-
nised nomenclaturally in this paper. In Western Australia is found a plant which is
superficially similar to the lectotype of subsp. inflata (from Queensland) but in
which the pericarp is almost glossy; to this variant belongs the lectoparatype of
Chenopodium inflatum collected by Morrison at the Ashburton River (PERTH).
This variant is further discussed under D. kalpari.

The characters which distinguished the two subspecies are far from constant and it
is evident that these taxa grade into each other. For this reason I have not kept them
as separate species as did both Aellen (1930) and Scott (1978b).

The plate in Mueller (1890 op. cit.) cited above under Chenopodium
rhadinostachyum appears to contain illustrations of two species; the small habit
drawing on the left-hand side is probably of D. kalpari while the drawing of the in-
florescence is of D. rhadinostachya. The illustrations of the seed incorrectly display
the position of the embryo and it is evident that both Mueller and Robert Graff, the
artist, had difficulty in establishing the orientation of this minute org an

The name Chenopodium rhadinostachyum var. paniculatum does not appear to
have been validly published. It was applied by Tate to a collection made by R. Helms
in 1891 on the Elder Exploring Expedition and, according to the text, was collected
at camp 53. 1 have seen only one specimen with that name; the label accompanying it
was written by Tate and the collection was made by Helms at camp 41. This speci-
men has a paniculate inflorescence typical of D. rhadinostachya but it is evident that
Tate assumed it to represent a new variety because specimens collected at camp 53
were identified by him as being the typical form of Chenopodium rhadinostachyum
whereas they are, in fact, the new species D. kalpari.

10. Dysphania kalpari Paul G. Wilson, sp. nov. Figure 7A.

Herba annua prostrata vel decumbens. Folia lamina profunde undulata, 1-5 cm
longa. Inflorescentia: spicae breviter pedunculatae, erectae, graciles, ad 15 cm longae.
Flores hermaphroditi et feminei; tepala 4, erecta obovata, c. 1 mm longa in base
connata, breviter hirsute; stamen 0-1; styli 2, graciles. Semen verticale compresse
sphaericum, c. 0.5 mm diam.; testa laevis, nitida; embryo semi-circularis basalis,
radicula cotyledonibusque superioribus.

Typus: West end of Hopkins Lake (south of Sir Frederick Range), Western Aus-
tralia, ± 24°20'S, 128°40'E; 1 Aug. 1962; D. E. Symon 2354 (holo: PERTH; iso: AD,
ADW).

Prostrate to decumbent annual herb with several stems arising from base, moder-
ately pilose all over with sessile or shortly stipitate glandular hairs and longer seg-
mented hairs, aromatic. Leaves thin, lamina elliptic, deeply undulate, 1-5 cm long;
petiole slender, half to as long as lamina. Inflorescence of axillary and terminal,
shortly pedunculate, erect, slender spikes to 15 cm long, made up of compact
dichasial cymes of 5-7 flowers subtended by bract. Bracts stipitate, limb ovate, ros-
trate, in all c. 2 mm long, not exceeding cyme. Flowers bisexual and female (the ter-
minal flower of cyme always bisexual); pedicel very short and thick, c. 0.2 mm long.
Tepals 4, erect, obovate, c. 1 mm long, united in lower third, rounded on back.

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Nuytsia Vol. 4, No. 2 (1983)

shortly hirsute; stamen 0-1, shortly exceeding perianth; styles 2, slender. Pericarp di-
aphanous (so thin as to he not apparent). Seed vertical, compressed spherical with
rounded margins (slightly keeled over embryo), c. 0.5 mm diam.; testa reddish brown
smooth and glossy; embryo semi-circular, basal, radicle and cotyledons superior,
raphe passing from upper margin to base. Infructescence breaking up when mature
with flowers dehiscing at apex of pedicel, the seed loosely enclosed in perianth.

QUEENSLAND: Boatman Stn., S. L. Everist 2935 (BRI); 7.8 mi [12.6 km] N of
Galway Downs, G. Trapnell E58 (BRD-

NEW SOUTH W.\LES: 70 km N of Bourke, S Jacobs 2061 (NSW); 8 mi [12.9 km] S
of Barrier Hwy on Cobar-Ivanhoe road, C. W. E. Moore 6012 (CANB).

SOUTH AUSTRALIA; 48 km W of Tallaringa Well, T. R. N. Lothian 3847 (AD); 30
mi [48.3 km] W of Welbourne Hill Stn., R. A. Perry 5534 (AD, MEL, NSW, NT,
PERTH).

WESTERN AUSTRALIA: Mundiwindi, B. G. Briggs 3601 (NSW); 44.5 mi [71.6 km]
E of Carnegie Stn., A. R. Fairall 1999 (PERTH).

NORTHERN TERRITORY; 16 mi [25.7 km] NW of Hamilton Downs, G.
Chippendale (NT 1779); 12 mi [19.3 km] W of Stuart Hwy on Yuendemu Rd, J. R.
Maconochie 60 (NT).

Distribution. Widespread in the inland areas of Australia in all mainland States ex-
cept Victoria. Map 22.

Habitat. Loam flats and red sand plains; often associated with mulga {Acacia
aneura).

Dysphania kalpari has been confused with D. rhadinostachya from which it may
be distinguished by its decumbent habit with several stems arising from the base, its
erect glossy seeds, and its “rats-tail” spikes. The habitat preferences of the two
species are quite different, those of D. rhadinostachya being skeletal soils often on
rocky slopes while D. kalpari is found on mulga {Acacia aneura) flats or sand plains.

Intergradation with Dysphania rhadinostachya subsp. inflata appears to occur, at
least with the variant of the latter found in the Pilbara region of Western Australia,
and it is possible that the lectoparatype of Chenopodium inflatum from the
Ashburton River area is a member of such an intergrade.

A note by F. T. Turvey with a collection made near Ernabella, South Australia, in
1968 (NSW 103747), states that “The natives gather the ripe seeds and make them
into a meal, and at this time mix them in with the honey from honey-ants (also in
season). This is relished as a food.”. On a specimen from Warburton Mission (R. A.
Gould, 3 April 1967, PERTH) it is recorded that the Aboriginal name is Kalpari and
that it is an important seed-food plant of ngatatjara, pitjantjatjara and pintupi
natives. A further specimen (W. H. Moyle, 1 July 1958, PERTH) has with it a note
to say that it is called Kalpari and that it is sought after by Aborigines in the Walter
James Range to make flour. The specific epithet is taken from the Aboriginal name
for the plant in central Australia.

Paul G. Wilson, Tribe Chenopodieae

197

3. SCLEROBLITUM Ulbrich

Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2, 16c: 495 (1934); Aellen in Hegi,
111. FI. Mitt.-Eur. ed. 2, 3: 660 (1960). Type: S. atriplicinum (F. Muell.) Ulbrich.

Chenopodium sect. Atriplicina Aellen, Verb. Naturf. Ges. Basel 41:99 (1931); A. J.
Scott, Bot. Jahrb. Syst. 100:211 (1978). Type: C. atriplicinum (F. Muell.) F. Muell.
(=S. atriplicinum).

Somewhat succulent very sparsely mealy herb. Stems numerous arising from root-
stock, unbranched. Leaves alternate simple, flat, the basal ones in a rosette. Flowers
in compact glomerules axillary to basal and stem leaves. Tepals 4, free; stamen soli-
tary in terminal flower, absent in others; ovary glabrous, stigmas 2, dorsiventral.
Pericarp hard and thick. Seed erect, completely filling utricle; embryo inverted hip-
pocrepiform (around apex of seed); endosperm sparse around embryo; perisperm
copious, central.

One species, endemic to Australia.

This genus is not closely related to any other Australian genus but appears to have
most in common with species of the genus Monolepis (northern hemisphere) and
Chenopodium sect. Thullingia (South America).

1. Scleroblitum atriplicinum (F. Muell.) Ulbrich in Engler et Prantl, Nat.
Pflanzenfam. ed. 2, 16c:495 (1934); Aellen in Hegi, loc. cit. — Blitum atriplicinum
F. Muell., Trans, et Proc. Viet. Inst. Advancem. Sci. for 1854-55: 133

(1855). — Chenopodium atriplicinum (F. Muell.) F. Muell., Fragm. 7:11 (1869);
F. Muell. Icon. Aust. Salsol. PI. t. 30 (1890). Lectotype: Cudnaka, Oct. 1850,
F. Mueller (MEL) lecto. nov.

Euxolus enervis F. Muell., Fragm. 1:140 (1859). — Amaranthus enervis (F. Muell.)
F. Muell. ex Benth., FI. Austral. 5:216 (1870). Type: In planitiebus ad flumina Darling
et Murray, J. Dallachy (holo: MEL).

Amaranthus tenuis Benth., FI. Austral. 5:216 (1870). Type citation: ‘Lower Darling
River, Herb. F. Mueller’ (iso: MEL).

Annual (or short-lived perennial?) somewhat succulent herb. Taproot prominent.
Stems numerous slender, prostrate or ascending, arising from rootstock, normally
unbranched. Leaves very sparsely mealy with thin-walled stipitate vesicular hairs,
eventually glabrous. Basal leaves in a rosette, lamina thin, 3-nerved, hastate-acumi-
nate c. 3 cm long with divaricate acuminate basal lobes; Petiole slender 3 to 8 cm long
with a broad clasping base; cauline leaves alternate, well separated, the upper ones
sub-sessile and narrowly elliptic. Flowers very shortly pedicellate; tepals 4, free,
erect, imbricate, ovate, acute, c. 1.5 mm long, somewhat glumaceous, mealy in lower
half with stipitate vesicular hairs; dorsal sepal larger than others and when mature
several nerved. Terminal flower of glomerule bisexual; stamen solitary opposite dor-
sal sepal, filament slender, glabrous, not thickened at base; anther caudate, c. 0.7 mm
long; ovary narrowly ellipsoidal, glabrous with a pair of opposite vascular traces
which pass up into the dorsi-ventral long slender stigmas. Ovule completely filling
ovary. Lateral flowers of glomerule female without staminodes. Fruiting perianth
cartilaginous; tepals narrowly oblong acuminate, c. 2.5 mm long, thickened keeled
and sigmoid at base. Utricle erect, lenticular; pericarp hard and thick; epicarp mem-
branous; mesocarp white hard and rugose, endocarp dark reddish brown, crus-
22493— (s)

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taceous, rugose on the outside. Seed erect, completely filling utricle and fused to the
endocarp; testa membranous, embryo inverted hippocrepiform (around apex of seed)
radicle inferior; perisperm central, copious, vitreous. Mature fruit shed with sur-
rounding perianth.

QUEENSLAND: Berbera, C. T. White 12231 (BRI); Clover Downs, R. Roe 3 (BRD-

NEW SOUTH WALES: Pinipara Ck., S. Jacobs 1003 (NSW); 3 mi [4.8 km] S of
Nyngan, G. M. Cunningham 766 (NSW).

VICTORIA; Gunbower, Oct. 1914, E. Curtis (NSW); Lake Hindmarsh, Oct. 1899
C. Walter (NSW.)

SOUTH AUSTRALIA: Yourambulla Caves, B. Copley 2241 (AD); 31 mi [49.9 km] N
of Overland Corner, D. E. Symon 3681 (ADW).

Distribution. Southern Queensland, central and southern New South Wales, north
west Victoria and eastern South Australia. Map 3.

Habitat. Heavy soils usually clay, frequently in situations that are seasonally water-
logged.

Although this plant has been referred to by collectors as an annual it has a sub-
stantial taproot which may enable it to live through more than one season.

The genus Scleroblitum was originally separated from Chenopodium largely be-
cause of the cartilaginous fruiting perianth; this in itself does not appear to be gen-
erically significant (especially as hardening of the tepals does not always take place),
however, the pericarp is very different from that in other Australian members of the
family. This pericarp has previously been interpreted as the testa and its distinctive
features overlooked: the developing seed fills the ovary at a very early stage causing
the testa to be in intimate contact with the ovary wall; this then presents difficulties
in distinguishing the two structures. The ovary wall is discernible, shortly after an-
thesis, as being composed of three layers; in the fruit these layers form very distinct
structures of the pericarp, an outer membranous epicarp in which a pair of faint vas-
cular strands pass up to the two style arms, a white crystalline mesocarp which be-
comes hard and rugose, and a dark reddish brown crustaceous endocarp. The endo-
carp is rugose on its outer surface (where in contact with the white mesocarp) and
smooth on the inner surface where it fuses with the membranous testa. The whole
structure is usually enclosed within the persistent cartilaginous perianth, however, in
some cases (possibly due to continuing moist conditions) the tepals remain small and
glumaceous.

Bentham (1870) described this species under three names, Chenopodium
atriplicinum, Amaranthus enervis and A. tenuis. The last two names were based on
material in which the fruiting perianth had not hardened. He also described the
plant (C. atriplicinum) as being slightly glandular-pubescent, a description repeated
by Black (1924, 1948) and by Scott (1978a). It was evidently for this reason that
Scott placed Chenopodium sect. Atriplicina in subgenus Ambrosia, along with the
glandular species of the genus, instead of in subgenus Chenopodium, in which are
placed the species with a mealy indumentum. In the latter subgenus, the sect.
Atriplicina comes closest to the sect. Thullingia in which the only species, C. antarc-
ticum (J. D. Hook.) J. D. Hook, is very similar, it also being a prostrate sparsely
mealy plant with a basal rosette of leaves, simple stems, glomerules of flowers with 2

Paul G. Wilson, Tribe Chenopodieae

199

to 4 tepals and solitary stamens, and erect seeds with a subannular embryo. The
genus Scleroblitum is, in addition, very similar in fruit and seed morphology to the
northern hemisphere genus Monolepis to which it is probably also closely related.

4. EINADIA Raf.

Rafinesque, Flora Telluriana 4:121 (1838); Dalla Torre et Harms, Gen. Siph. 145
(1900), pro syn. sub Suaeda; Ulbrich in Engler et Prantl, Nat. Pflanzenfam. ed. 2,
16c: 555, 558 (1934) pro syn. sub Suaeda sect. Schanginia\ A. J. Scott, Feddes Repert.
80:3 (1978). Type: Einadia linifoUa (R.Br.) Raf.

Chenopodium sect. Polygonoidea Aellen, Feddes Repert. 69:69 (1964) et in Hegi, 111.
FI. Mitt.-Eur. Ed. 2, 3/2:577(1960) sine desc. latina. Type: Chenopodium poly-
gonoides (Murr) Aellen.

Herbaceous or weakly woody perennials, often prostrate or scandent, branching
from the base. Branches slender, ridged. Leaves opposite or alternate, petiolate, lin-
ear to broadly hastate, c. 2-3 cm long, flat and slightly fleshy, ± covered when young
(at least below) with spherical vesicular hairs. Flowers small, gynomonoecious in
small cymose clusters forming racemose or open panicles, often congested; pedicels
(when present) constricted at apex. Terminal flower of cluster bisexual; lateral
flowers female without staminodes. Tepals (4) 5, herbaceous, sometimes succulent in
fruit. Stamens 1-3, often spreading at anthesis with the subtending tepal, ± equal to
tepals, glabrous; filament slightly pulvinoid at base; disc absent. Ovary glabrous;
style absent or very short; stigmas 2, slender, papillose. Fruit a nut with thin dry
pericarp or depressed globose (rarely ovoid) berry with inflated pericarp (succulent
outside, water within), not enveloped by perianth. Seed horizontal, depre.ssed globose
to lenticular with rounded margin; testa crustaceous; embryo annular, radicle
centrifugal; perisperm central and copious.

A genus of six species endemic to Australia and New Zealand.

Rafinesque (1838 op. cit.), in his original description, segregated the genus Einadia
from Rhagodia on the basis of the presence of only one or two stamens instead of
five. In this he was relying solely on Robert Brown’s descriptions of the seven species
of Rhagodia since he had seen no material. Brown (1810) only in R. linifolia referred
to the stamen number (as being 1 or 2), evidently he assumed it to be 5 in the other
species as indicated in his generic description. Rafinesque, therefore, only
transferred this one species to Einadia and was unaware of its close relationship (and
common stamen number) with R. nutans and R. hastata which had also been de-
scribed by Brown. The generic name Einadia was subsequently ignored or misap-
plied until Scott (1978a) took it up. Although Scott recognised the marked difference
between those species of Rhagodia with polygamous flowers and reduced stamen
number (which he placed in Einadia) and those with dioecious flowers and five
st am ens (which he retained in Rhagodia), he did not comprehend the close relation-
ship of the Einadia group with those Australian and New Zealand species of Cheno-
podium that have similar floral characters and similar habit but which lack a succu-
lent pericarp. The similarity is such that with non-fruiting material it is often im-
possible to recognise to which genus it should be assigned. The difficulty is enhanced
by the fact that the succulent pericarp does not always develop in the group of
species with normally baccate fruits and, further, that hybridization appears to take
place between related species placed in the different genera. I am therefore ex-
panding the genus Einadia to include those Australian and New Zealand species of

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Chenopodium and Rhagodia with a herbaceous perennial habit, small flat leaves
with discrete vesicular indumentum and polygamous 5-merous flowers with reduced
stamen number.

The distinction between Einadia and Chenopodium is not easy to define, partly
because of the great variability found in the latter genus. Einadia comes closest to a
group of species in Chenopodium sect. Leprophyllum (e.g. C. hubbardii) and another
in sect. Desertorum (e.g. C. desertorum). From the former section the species of
Einadia differ in being perennials (not annuals) and in having 1-3 (not 5) stamens,
while from the latter section they differ in absence of disc (pubescent disc in sect.
Desertorum) and again reduced stamen number.

The species of Einadia differ from those of Rhagodia sensu stricto in being her-
baceous usually sprawling perennials with gynomonoecious flowers, a reduced
stamen number and no disc. The fruit, if succulent, is “inflated”, that is it has a
fleshy outer and a watery inner layer; or drying it does not become viscous. The in-
dumentum is of spherical hairs (which may collapse to become saucer — or shallowly
cup-shaped) and not of irregularly shaped hairs, nor does it collapse to form a scaly
sheen as it does in some Rhagodia species.

It is not clear why Dalla Torre et Harms (1900 op. cit.) made Einadia a synonym of
Suaeda-, possibly they assumed, as did Ulbrich (1934 op. cit.), that Einadia linifolia
was a synonym of Suaeda linifolia Pallas, which is not the case; the two species are
very distinct as are their respective genera.

Key to species

1. Fruit succulent 2

Fruit dry

2. Leaves deltoid or broadly hastate with rounded apex, dark green . . 5. E. hastata
Leaves linear to ovate, deltoid, narrowly hastate or sagittate, acute or obtuse.

grey to green 3

3. Leaves ovate, obtuse, base usually sagittate (N.Z.) 2. E. triandra

Leaves linear to broadly triangular, base often hastate 1. E. nutans

4. Tepals in fruit with a hard circular (often black) limb; leaves narrowly oblong-

elliptic to narrowly hastate 6. E. polygonoides

I epals in fruit oblong to obovate, herbaceous or indurated; leaves broadly ovate
or elliptic to orbicular or deltoid 5

5. Leaves broadly elliptic to orbicular (N.Z.) 4. E. allanii

Leaves broadly ovate to deltoid, often hastate at base 3. E. t'rigonos

1. Einadia nutans (R.Br.) A. J. Scott, Feddes Repert. 89:3 (1978).— Rhagodia
^tans R.Br., Prod. 408 (1810). Lectotype. Ad rupes saxa in Oyster Cove in fluvio
Derwent, Feb. 1804, R. Brown (holo: BM) lecto. nov. Figure 4E.

R. linifolia R.Br., Prod. 408 (1810). — E. linifolia (R.Br.) Raf., FI. Tellur. 4:121 (1838).
Lectotype: Broad Sound, Sept. 1802, R. Brown (holo: BM; iso: K, NSW) lecto. nov.

R. chenopodioides Moq., Chenop. Monogr. Enum. 11 (1840). Lectotype: Port
Jackson, 1830, Gaudichaud (holo: P) lecto. nov.

Chenopodium australasicum Moq., Chenop. Monogr. Enum. 20 (1840) Type ci-
tation: “Nova Hollandia, Caley’\ Type: P, n.v.

Paul G, Wilson, Tribe Chenopodieae

201

Scrambling perennial with woody base. Branches slender, ridged, sparsely covered
when young with small discrete bladder hairs. Leaves opposite or alternate, lamina
chartaceous to sub-coriaceous, linear to elliptic or narrowly to broadly triangular, en-
tire 10-20(40) mm long, rounded to truncate or hastate at base, sparsely to moder-
ately mealy when young, eventually glabrescent (at least above), abruptly petiolate.
Inflorescence a narrow racemose panicle 2-4 cm long; flowers in glomerules, either
compact or pedicellate, if terminal then bisexual, otherwise usually female. Perianth
depressed globular, 0.7-1.5 mm diam., not enveloping the ovary, mealy with bladder
hairs to glabrescent; tepals (4)5, oblong, somewhat enlarging and becoming red,
fleshy and sometimes reflexed in fruit. Bisexual flower: stamens 1-2(3), spreading or
erect at anthesis; filament ligulate, thickened at base, glabrous; anthers included, c.
0.3 mm long; disc absent; ovary ovoid, glabrous, stigma 2 sessile or subsessile, divari-
cate, terete, 0. 1-0.2 mm long. Female flower: staminodes absent; ovary as in bisexual
flower. Fruit: pericarp inflated succulent, orange or red, depressed (or narrowly
ovoid), c. 4 mm diam., exceeding perianth, surface faintly colliculate; seed lenticular
with rounded margin, 1-1.5 mm diam.; testa rugulose to areolate-reticulate, black.

Distribution. Central and eastern Australia including Tasmania.

This species exhibits considerable variation with several of the variants having
broad regional distributions. In many cases these variants are partially sympatric
although evidently ecologically separated. Transition between the variants is readily
apparent from study of the herbarium material. Only four of the variants have been
recognised in this paper leaving a large number of specimens which it is only possible
to determine to the species level. To attempt a more complete taxonomic analysis
would appear to be unwarranted due to the considerable plasticity of the plants, and
their transition in the field from one variant to another. Vegetative or non-fruiting
material is sometimes difficult to distinguish from E. trigonos subsp. trigonos and
specimens of the two species were confused by Bentham (1867) and by later authors;
it seems likely, from the observation of herbarium material, that the two hybridize
but the study of field evidence is required to support this. Hybridization between E.
nutans and E. hastata may also occur since several collections appear to be
intermediate in morphology.

The name Rhagodia nutans was applied for some time to the New Zealand plant
now known as Einadia triandra (e.g. by Cheeseman 1884 and 1914). The latter
species is very similar to some variants of E. nutans and, if native to Australia, would
probably be included with it under the earlier name E. triandra. One of the variants
of E. nutans found in southern Australia is so similar to the New Zealand plant as to
make identification somewhat dependent on locality data. Einadia nutans subsp.
nutans is found in New Zealand as a recent introduction and is there very distinct in
appearance and ecology from the native E. triandra. It forms dense hummocks or
scrambles over hedgerows so as to kill the surrounded shrubs (A. J. Healy in sched.
CHR 91541). The characters by which E. nutans may be distinguished from E.
trigonos are discussed under the latter species.

The name Rhagodia chenopodioides Moq. was considered by Bentham (1870), who
had not seen the type, to be possibly a synonym of R. billardieri (= R. baccata) and
was included by Scott (1978a) under this latter name. The type is, however, a nar-
row-leaved varient of Einadia nutans subsp. nutans.

The terminal flower of a branch of the inflorescence is normally bisexual but may
be functionally male through the weak production of stigmas. In most specimens
studied this flower had normal stigmas and produced a seed.

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Key to subspecies

1. Leaves linear to narrow oblong Ic. subsp. linifolia

Leaves either broader than above or hastate to sagittate at base 2

2. Leaves narrowly sagittate, sub-coriaceous; seed c. 1.5 mm diameter (central

Australia) Id. subsp. eremaea

Leaves narrowly to broadly hastate or deltoid, thin; seed c. 1 mm diameter ... 3

3. Leaves narrowly hastate; berry ovoid lb. subsp. oxycarpa

Leaves variable in shape; berry depressed globular (if ovoid then leaves
deltoid-hastate) la. subsp. nutans

la. subsp. nutans

R. nutans var. parvifolia Moq. in DC., Prod. 13/2:54(1849). Lectotype: Ex herb. Fee
(G-DC, photo seen) lecto. nov.

Chenopodium triangulare subsp. convolvulinum Murr, Allg. Bot. Z. Syst. 16:56
(1910).— C. triangulare var. convolvulinum (Murr) Maiden, Census N.S.W. Plants 66
(1916). Syntypes: Stonehenge, Dec. 1899, J. H. Maiden (NSW 33761); Jenolan Caves,
Dec. 1899, W. F. Blakely (NSW 142922). Lectotype: Stonehenge, Dec. 1899, J. H.
Maiden (NSW 33761) Lecto. nov.

? R. nutans var. fallacina Domin, Biblioth. Bot. 89: 616 (1921). Type: Cheltenham,
Victoria, Domin IV. 1910, n.v.

R. chenopodioides Moq., op. cit. (1840).

Leaves chartaceous, hastate to deltoid and shortly lobed at base. Inflorescence
slender or branched near base; fruits sessile or pedicellate. Berry depressed globular;
seed variably reticulate, c. 1 mm diameter.

QUEENSLAND: Tballon, R. Roe 26 (CANB); 10 mi [16.1 km] N of Wondoan, N. H.
Speck 1961 (NSW).

NEW SOUTH WALES: 5 mi [8 km] S of Cobar, C. W.E. Moore 4480 (CANB);
Brush Island, 5 Sept. 1937, F. A. Rodway (NSW).

VICTORIA: 11.6 km from Dargo, E. M. Canning 3937 (CBG); Nandaly, W. W. Watts
465 (NSW).

TASMANIA: Lewisham, 3 Mar. 1952, W. M. Curtis (HO); Carlton Bluff, 19 Feb. 1959,
W. D. Jackson (HO).

SOUTH AUSTRALIA: Vihlpena Pound, A. E. Orchard 2594 (AD); Oraparinna Nat.
Park, J. Z. Weber 2578 (AD).

NORTHERN TERRITORY: Mt Le Hunte, P. K. Latz 913 (NT); 30 km S of Alice
Springs, R. Pullen 10556 (CANB).

Distribution. Central and south-eastern Australia including Tasmania. Map 23.

Habitat. Occupying a wide range of habitats but usually in loam or heavy soil.

This polymorphic subspecies comprises, apart from the type variant, those vari-
ants not included in the other three subspecies. In New South Wales it grades to the
north into subsp. linifolia and to the north-west into subsp. eremaea. However,
plants with the leaf shape of subsp. nutans are also found in northern New South

Paul G. Wilson, Tribe Chenopodieae

203

Wales and southern Queensland where plants intermediate between these and subsp.
linifolia also occur. A variant of subspecies nutans very similar in appearance to the
east coast plant also occurs in the southern portion of the Northern Territory and in
adjacent regions of South Australia; it is here sympatric with subsp. eremaea
although probably ecologically isolated being an inhabitant of the ranges rather than
the sandplains. No putative hybrids between the two subspecies in this area have
been seen.

A variant of Einadia nutans found in south-east Queensland appears to differ
significantly from the other variants referred to subsp. nutans. It is a more delicate
plant than typical subsp. nutans with very slender branches. The leaves are predomi-
nantly opposite, thin, with the lamina bluntly deltoid-hastate, and c. 10 mm long in
the south-east becoming larger (c. 15 mm long) farther north. The inflorescence is
slender with fruits on long slender pedicels c. 5 mm long. The tepals enlarge so as
sometimes to almost cover the young fruit but eventually are reflexed. The berry in
some specimens appears to be ovoid in shape as in subsp. oxycarpa but mature fruit
is not usually present and no collector has commented on its appearance. The follow-
ing specimens are examples of this ‘delicate’ variant in eastern Queensland: 2 mi [3.2
km] S of Inverai School, R. W. Johnson 432, 433 (BRI); Bybera, C. T. White 10387,
10405 (BRI); Mungallala, C. E. Hubbard 6025 (BRI); Hannaford, L. Pedley (BRI,
CANB); Chinchilla, A. Beasley 34 (BRI); Woodlands, T. J. McDonald 196 (BRI);
Coppersmith Rock, G. Tandy (BRI).

lb. subsp. oxycarpa (Gauba) Paul G. Wilson stat. et comb. nov. — Rhagodia
nutans var. oxycarpa Gauba, Viet. Nat. 65:167 (1948). Type: Near Loveday, South
Australia, 28 May 1943, E. Gauba (holo: MEL 80685).

Scrambling sub-sbrub to 1 m high. Leaves somewhat leathery, narrowly hastate
acute, c. 15 mm long, lateral lobes narrow, acute. Inflorescence branched; flowers and
fruits sessile. Fruiting perianth fleshy and reflexed, red; berry ovoid, c. 4 mm long,
pericarp colliculate; seed horizontal to erect c. 1 mm diameter, testa moderately re-
ticulate.

NEW SOUTH WALES: 56 km N of Wentworth, 29 Aug. 1969, A. Rodd (NSW); 10
km ESE of Mungindi, A. Rodd 1070 (NSW).

SOUTH AUSTRALIA: 2 mi [3.2 km] NW of Lake Bonney, B. Copley 2577 (AD);
Koonamore, M. Crisp 565 (AD).

Distribution. New South Wales and South Australia. Map 24.

Habitat. Often on sandhills but also recorded in other situations.

This subspecies cannot be readily distinguished when in flower from some variants
of the polymorphic subspecies nutans; it usually has thicker and narrower leaves
which have narrow divaricate basal lobes, however, since leaf characters are
notoriously plastic in this genus it is really only by the ovoid fruit that subsp.
oxycarpa may be positively identified.

According to M. D. Crisp (pers. comm.) subsp. oxycarpa is edaphically segregated
from subsp. nutans, the former being found on the top of sand dunes and the latter
on the flats. While this is evidently the case in some areas, over its total range subsp.
oxycarpa occupies a variety of habitats.

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The wide and scattered distribution of subsp. oxycarpa suggests that it may rep-
resent a fruit-form which appears sporadically in populations of subsp. nutans, but
extensive field and experimental work is required to establish the biological situ-
ation.

Ic. subsp. linifolia (R. Br.) Paul G. Wilson, Comb, et stat. nov. — Rhagodia
linifolia R.Br., Prod. 408 (1810). — Einadia linifolia (R.Br.) Raf. l.c. (1838); Ulbrich in
Engler et Prantl. Nat. Pflanzenfam. ed. 2, 16c; 558 (1934) pro syn. sub Suaeda
linifolia Pallas. — E. nutans var. linifolia (R.Br.) A. J. Scott, Feddes Repert. 89:4
(1978). Lectotype: see above.

Chenopodium australasicum Moq., op. cit. (1840), see note below.

Prostrate or scrambling perennial. Leaves thin, linear to narrow-oblong.
Inflorescence slender; fruits frequently slender pedicellate. Tepals enlarged and
spreading or reflexed in fruit; berry depressed globular; seed finely reticulate, 1 mm
diameter.

QUEENSLAND; Noondoo Stn, S. L. Everist 829 (BRI); East of Condamine, W. T.
Jones 3733 (CANB).

NEW SOUTH WALES; Warrah, Mar. 1920, W. W. Froggatt (BRI); 5 mi [8 km] from
Aberdeen, R. Story 1092 (CANB).

VICTORIA; Echuca, H. King 4 (MEL).

Distribution. Eastern Queensland and New South Wales, northern Victoria. Map 24.

Habitat. Predominantly found on heavy soils.

The typical variant of this subspecies is found in north east Queensland and east-
ern New South Wales. In New South Wales it grades into subsp. nutans and to the
west into subsp. eremaea. Some inland variants have rather thick leaves and com-
pact inflorescences, possibly as a result of soil salinity, but this is not evident from
the label data on the herbarium specimens.

I have not seen the type of Chenopodium australasicum, however, Moquin’s de-
scription corresponds to that of subsp. linifolia and it was with this taxon that he
later (1849) synonymized the name.

A variant of Einadia nutans with very narrow leaves is found in N.W. Victoria and
south central New South Wales but the material I have seen is too incomplete to de-
termine its precise affinities. I have included this variant within subsp. linifolia on
the basis of its leaf shape.

The synonymizing of E. linifolia under Suaeda linifolia Pallas by Ulbrich (1934)
was presumably made on the assumption that they were nomenclatural synonyms.
This is not the case and I do not know why Ulbrich came to this conclusion.

Id. subsp. eremaea Paul G. Wilson, subsp. nov.

Herba perennis leviter scandens. Rami graciles sulcati. Folia opposita vel alterna,
cinerea; lamina aliquantum coriacea, anguste triangularis ad basim sagittata, 15-
40 mm longa. Flores fructusque sessiles. Baccus depresso globularis. Semen c. 1.5 mm
diam.; testa valde alveolata.

Paul G. Wilson, Tribe Chenopodieae

205

Typus: Close to Ayers Rock, Northern Territory, sandy soil, 25 Aug. 1973, A. A.
Munir 5132 (holo: AD 97342352; iso: DNA, NT).

Scrambling perennial herb to 1 m high with slender sulcate branches. Leaves op-
posite or alternate, grey; lamina somewhat coriaceous, narrowly triangular, sagittate
at base, 15-40 mm long. Inflorescence paniculate, branched near base. Flowers and
fruit sessile. Berry depressed globular; seed c. 1.5 mm diameter, testa strongly alveo-
late.

QUEENSLAND: Earlstoun Stn., S. T. Blake 5480 (BRI); Boorara, Vk. J. Bisset 253
(BRI).

NEW SOUTH WALES: Tundulya, C. W. E. Moore 4559 (CANB).

SOUTH AUSTRALIA: 35 km W of Mabel Creek, T. R. N. Lothian 2783 (AD); 48 km
W of Todmorden Stn., T. Reichstein 24 (AD).

WESTERN AUSTRALIA: 8 km S of Giles, R. H. Kuchel 161 (AD); Giles Settle-
ment, R. Hill 1369 (AD).

NORTHERN TERRITORY: Tanami Sanctuary, T. S. Henshall 1256 (NT);
Petermann Creek, N. M. Henry 576 (NT).

Distribution. Central and Western Queensland, north western New South Wales,
northern South Australia, east-central Western Australia, the Northern Territory.
Map 25.

Habitat. Generally in sand.

This subspecies is sympatric with subspecies nutans in parts of the Northern Ter-
ritory and in the Musgrave Ranges area of South Australia, where the two evidently
occupy different ecological niches (subsp. eremaea in sand and subsp. nutans on hilly
country) and I have seen no suggestion of intergradation between the two in that re-
gion. Subspecies eremaea is a more robust plant than any of the other subspecies and
is bigger in all its parts; it grades to the east into subsp. linifolia and to the south and
south-east into subsp. nutans.

2. Einadia triandra (G. Forster) A. J. Scott, Feddes Repert. 89:5
(1978). — Chenopodium triandrum G. Forster, FI. Ins. Austr. 21 (1786); Moq., Chen.
Mon. Enum. 25(1840) p. p.; J. D. Hook., FI. Nov.-Zel. 1:212 (1857) p.p.; J. D. Hook.,
Handb. N. Zeal. FI. 230 (1864) p.p. — Rhagodia triandra (G. Forster) Aellen,
Candollea 8:5 (1939); Allan, FI. N. Zeal. 1:231 (1961). Type: Queen Charlotte Sound,
New Zealand, J. R. and J. G. A. Forster (holo: BM; iso: K n.v. fide Allan 1961). Fig-
ure 5B.

[Rhagodia nutans auct. non R.Br.: Cheeseman, Man. N. Zeal. FI. ed. 2, 404 (1925)]

Spreading or scrambing perennial with woody (sometimes rhizomatous?) rootstock
and numerous slender matted stems. Branches striate, mealy when young. Leaves
opposite or alternate, mealy or glabrescent; lamina thin, ovate, 5-10 mm long, entire,
apex obtuse and minutely apiculate, base truncate to shortly sagittate; petiole slen-
der c. 4 mm long. Inflorescence of glomerules in terminal or axillary slender racemose
panicles c. 5 cm long in fruit. Flowers globular 1-2 mm diam.; tepals ovate, flat or cu-
cullate, enlarging and becoming fleshy in fruit, mealy outside; stamens 0-3, spreading

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Nuytsia Vol. 4, No. 2 (1983)

at anthesis; filament thickened at base; stigmas 2 sessile, slender c. 0.5 mm long.
Fruit a berry; pericarp red, globular, succulent c. 4 mm diam. Seed depressed sphe-
roid, c. 1.7 mm diam, testa prominently reticulate.

NEW ZEALAND: N of Kaikoura, R. Melville 5833 (CHR); Near Cape Campbell, W.
R. Sykes 500/70 (CHR).

Distribution. New Zealand; ‘from near north Cape to a little south of lat. 44°’ fide
Allan 1961.

Habitat. ‘Rocky and gravelly coastal places’ fide Allan 1961.

I have not seen the type of Einadia triandra but the description of the leaves given
by J. G. Forster agrees with current concepts and with the plant described under the
name Rhagodia triandra by Aellen (1939).

The transfer of Chenopodium triandrum to Rhagodia was intimated by F. Mueller
in 1886 and 1887 when he wrote ‘The Chenopodium triandrum of New Zealand is
transferable to the genus Rhagodia', but I cannot find any publication in which he
made the combination.

The name Chenopodium triandrum G. Forster was considered by Murr (1910) to
include C. microphyllum F. Muell., and his synonymy was accepted by Maiden and
Betche (1916). The two species are, in fact, quite distinct and differ, among other
characters, in stamen number and fruit type, although it is probable that Murr had
specimens of C. allanii ( = Einadia allanii) in mind rather than C. triandrum when he
suggested the synonymy.

Einadia triandra is very similar to some variants of E. nutans and if it were not for
the fact that the one is endemic to New Zealand and the other to Australia a
subspecific status for E. nutans would probably be more suitable. The leaves of E.
triandra have consistently an ovate shape obtuse at apex and truncate at base
usually with short downwardly directed lobes (sagittate) whereas in E. nutans the
leaves tend to be triangular, acute at apex and hastate at base (i.e. with divaricate
lobes). In E. nutans the seed is lenticular with rounded margin and the surface
rugulose to deeply reticulate; in E. triandra the seed is depressed-spheroid (i.e.
thicker than in E. nutans) and consistently deeply reticulate.

Prior to the paper by Aellen (1939) material of Einadia triandra had usually been
referred to Rhagodia nutans and the name Chenopodium triandrum incorrectly ap-
plied to the species described by Aellen as C. allanii. The last species may be dis-
tinguished by its thin dry pericarp; it also differs from all related species found in
New Zealand by its entire, elliptic to orbicular leaves.

The recording by Sykes (1977) of E. triandra (as Rhagodia triandra) in the
Kermadec Islands was based on several collections some of which are of a variant of
E. trigonos, q.v. while others are of an undescribed species.

3. Einadia trigonos (Roemer et Schultes) Paul G. Wilson, comb.
nov.— Chenopodium trigonon Roemer et Schultes, Syst. Veg. 6:275 (1820) based on
following.— C. triangulare R.Br., Prod. 407 (1810) nom. illeg. non Forsskal (1775).
Type: Nepean and Hawkesbury Rivers, R. Brown (holo: BM).

C. stellulatum Aellen in Probst, Mitt. Naturf. Ges. Solotb. 8:57 (1928). Type:
Wollkompost der Kammgarnfabrik Derendingen, 1916, R. Probst, n.v.

Paul G. Wilson, Tribe Chenopodieae

207

Weak prostrate to erect straggling perennial herb branching from the base.
Branches slender, faintly striate, glabrous or sparsely mealy when young. Leaves
alternate or subopposite, thin, lamina broadly ovate to deltoid (or hastate with short
spreading lobes), 8-15 mm long, glabrescent above, sparsely to moderately mealy
below with white spherical vesicles; petiole slender, short or equal in length to
lamina. Inflorescence a congested to slender terminal and axillary panicle with
numerous sessile flowers aggregated into glomerules. Flowers, if terminal to a partial
cluster, bisexual; the others female. Perianth depressed globular 0.5-1 mm diam.;
tepals broadly ovate to obovate, spathulate or linear, often cucullate, sparsely mealy
or glabrous. Bisexual flower, stamen solitary, divaricate at an thesis, filament
thickened (pulvinoid) at base; ovary depressed, papillose; stigmas slender, spreading,
± sessile. Female flower: stamens and staminodes absent, otherwise as in bisexual
flower. Fruit: tepals narrow to broad oblong or obovate, herbaceous or indurated,
smooth, if indurated then with a slender claw and cucullate limb enclosing and per-
sisting around fruit; pericarp dry (?slightly succulent in subsp. leiocarpa), thin, often
black and usually prominently papillose. Seed lenticular with rounded margin, c.
1 mm diam.; testa slightly granular (or striate in subsp. leiocarpa).

In habit and leaf shape E. trigonos is similar to some variants of E. nutans but in
the latter species the leaves are usually more prominently hastate. The tepals of E.
trigonos vary from being broad-oblong to linear. The variant with broad tepals
(in subsp. trigonos) when in flower is difficult to distinguish from some variants of E.
nutans and even fruiting material can be troublesome since, although the latter
species usually has baccate fruit, the succulence does not always develop or is not ap-
parent in dried material. A further distinction is found in the seed which in E.
trigonos is granular and the pericarp smooth to strongly papillate (but see also
subsp. leiocarpa); in E. nutans the seed is alveolate to rugulose and the pericarp
smooth or colliculate (i.e. with low convex markings). It appears probable, on the
basis of herbarium material, that the two species do hybridize.

To separate Einadia trigonos from E. hastata is also sometimes difficult in the ab-
sence of fruit, however, the leaves of E. hastata are usually darker green, thicker, and
they are nearly always opposite, whereas in E. trigonos they are predominantly alter-
nate.

Key to subspecies

1. Tepals spathulate to linear; pericarp strongly papillose, frequently black

3b. subsp. stellulata

Tepals ovate or elliptic to obovate; pericarp almost smooth 2

2. Leaves broadly hastate or deltoid with rounded apex 3a. subsp. trigonos

Leaves hastate to narrow hastate, acute 3c. subsp. leiocarpa

3a. subsp. trigonos

Plant prostrate to decumbent. Leaves broadly hastate or deltoid with rounded
apex. Tepals ovate to obovate, herbaceous in fruit. Pericarp almost smooth.

NEW SOUTH WALES: Moona Plains, Feb 1899, S. R. Crawford (NSW); Conjala,
May 1899, W. Heron (NSW).

VICTORIA; Mt Tingaringy, A. C. Beauglehole 35726 (MEL); Melbourne, 1886, J.
Minchin (MEL).

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Nuytsia Vol. 4, No. 2 (1983)

NEW ZEALAND: Coppermine Is., Mauraki Gulf, 25 Oct. 1968, M. A. and I. M.
Ritchie (CHR); Little Sister Is., 25 Oct. 1973, B. D. Bell (CHR); Curtis Island,
Kermadec Group, W. R. Sykes 923/K (CHR).

Distribution. Southern Victoria and extreme eastern New South Wales to just north
of Walcha. New Zealand (an atypical variant). Map 26.

Habitat. Often near water but also in open country or forested areas.

The typical variant of this subspecies has thin, deltoid to broadly hastate leaves
and broadly ovate tepals, it is found in the extreme south east of Victoria north to an
area just north of Sydney. Farther north still the tepals become narrower and the
plant grades into subsp. stellulata which has linear tepals.

The presence of Einadia trigonos subsp. trigonos in New Zealand is probably due
to its relatively recent introduction. The earliest collection seen by me is dated 1926
although only the specimens in herb. CHR have been studied. In New Zealand it is
variable in leaf form and whereas some of these variants are found in Australia
others are distinct and approach in aspect, and in flowers and fruit, E. allanii, a
species endemic to New Zealand, suggesting that hybridization may be taking place.
Sykes (1977) recorded Rhagodia triandra from several islands in the Kermadec Is.
group but the specimens I have seen from these islands which have been so anno-
tated either belong to the New Zealand variant of E. trigonos subsp. trigonos or to an
undescribed species. Sykes also recorded Einadia allanii (as Chenopodium allanii),
based on the presence of its seed in the crop of a parakeet; since, however, the seed
of E. allanii is virtually indistinguishable from that of E. trigonos it seems likely that
they came from plants of the latter species which on some islands was recorded as
abundant.

The New Zealand specimens of subsp. trigonos have flowers with broad tepals
which enlarge with the fruit and almost envelop it at maturity; many also have pur-
plish stems and leaf margins, characters not observed (or not noted) on Australian
specimens.

3b. subsp. stellulata (Benth.) Paul G. Wilson, comb, et stat. nov. — Chenopodium
triangulare var. stellulatum Benth., FI. Austral. 5:161 (1870).— C. stellulatum
(Benth.) Aellen, Verh. Naturf. Ges. Basel 41:93 (1931) nom. illeg. non Aellen (1928).
Type citation: “New England, C. Stuart.” Lectotype: River Severn. C. Stuart 44
(MEL 80677) lecto. nov.

C. stellulatum Aellen in Probst, op. cit.

Prostrate to spreading herb with stout taproot. Inflorescence congested when in
flower, elongating in fruit to a slender racemose panicle with disjunct glomerules of
flowers. Tepals spathulate to linear, frequently indurated in fruit. Pericarp strongly
papillose, frequently black.

QUEENSLAND: Waterford, W. Jones 3345 (CANB); Cooranga North, C. T. White
2492 (BRI).

NEW SOUTH WALES: Grafton, H. Henholz (NSW); Swan Bay, 8 June 1957,
L. A. S. Johnson (NSW).

NEW ZEALAND: Tai Tapu, Canterbury, A. J. Healy 54/123 (CHR); Between
Halswell and Tai Tapu, 24 Aug. 1949, S. E. Archer (CHR 68003).

Paul G. Wilson, Tribe Chenopodieae

209

Distribution. South-east Queensland and eastern New South Wales. A recent
introduction in New Zealand. Map 27.

Habitat. In partial shade frequently in moist situations in heavy soil. A ready colon-
iser of disturbed situations.

The vernacular names of Tar vine and Fish weed applied to this plant are a
reflection of the strong fish smell possessed by some variants; it has been reported to
cause cows to produce tainted milk (C. T. White 1921 as Chenopodium triangulare).

As a recent introduction it is stated to be well established along a roadside near
Halswell in the South Island of New Zealand (A. J. Healy 1954 in sched.)

3c. subsp. leiocarpa Paul G. Wilson, subsp. nov.

Herba perennis erecta, ad basim ramificans. Folia alterna; lamina hastata vel
anguste hastata acuta, c. 15 mm longa. Pericarpium laeve, translucens (non
denigratum). Semen c. 1.2 mm diam., testa striata.

Typus\ Pilliga, New South Wales, Nov. 1932, H. M. R. Rupp 5 (holo: NSW 142708).

Erect perennial herb branching from base. Leaves alternate; lamina hastate to nar-
row hastate (upper ones narrow elliptic), c. 15 mm long, acute, noticeably mealy
below; petiole slender, c. half length of lamina. Tepals broadly elliptic to obovate.
Pericarp smooth, translucent (not blackened), evidently herbaceous when mature.
Seed c. 1.2 mm diam.; testa glossy, radially striate.

NEW SOUTH WALES: Gurley, Nov. 1914, E. Breakwell (NSW); Boggabri, R. H.
Cambage 3655 (NSW); 6.5 mi [10.5 km] W of Wee Waa, 19 Dec. 1966, L. A. S.
Johnson (NSW); Moree, Feb. 1907, J. T. Scott (NSW).

Distribution. North-east New South Wales. Map 26.

Habitat. Only recorded with one collection as “Common on heavy grey alluvium
under Eucalyptus camaldulensis”.

This subspecies is distinctive in having small narrowly hastate leaves and in
having a fruit with a smooth transluscent pericarp; the striate seed also clearly separ-
ates it from the other subspecies. The subspecies leiocarpa is possibly more closely
related to E. nutans, with which the seed ornamentation and herbaceous pericarp
would have more in common, but it seems preferable to restrict the latter species to
those plants with a succulent fruit.

A collection from 16 mi [25.7 km] W of Garah (K. L. Soiling 431, NSW) differs
from typical subsp. leiocarpa in having a more triangular leaf, a thin pericarp and an
almost smooth seed. From the same locality has been collected a plant with leaf
shape typical of subsp. leiocarpa but with tepals in fruit somewhat succulent and a
pericarp thin but succulent (K. L. Soiling s.n. NSW 142982). It appears to be
intermediate in morphology between E. trigonos subsp. leiocarpa and E. nutans.

The infraspecific epithet is derived from two Greek words, leios, smooth, and
carpos, fruit, referring to the smooth pericarp.

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4. Einadia allanii (Aellen) Paul G. Wilson, comb nov. —Chenopodium allanii
Aellen, Candollea 8:7 (1939). Lectotype: Lammermoor Mts, New Zealand, Jan. 1929,
H. H. Allan (CHR 1064) lecto. nov. Figure 5A.

[Chenopodium triandrum auct. non G. Forster: J. D. Hook., FI. Nov.-Zel. 1:212
(1853) p.p.; J. D. Hook., Handb. N. Zeal. FI. 230 (1864) p.p.; ? T. F. Cheeseman, 111.

N. Zeal. FI. 2:t.l64 (1914).]

Spreading perennial woody towards base. Branches weak, slender, glabrescent.
Leaves alternate; lamina thin, broadly elliptic to orbicular, 5-15 mm long, entire,
sparsely mealy to glabrous; petiole slender. Inflorescence of axillary and terminal
slender or branched panicles (congested when young). Flowers c. 0.5 mm diameter;
tepals (4)5, ovate, not cucullate, united towards base, moderately mealy outside, en-
larging in fruit but not enclosing nut; stamens 0-3, slightly exceeding tepals; fila-
ments thickened at base, glabrous, disc absent; style absent, stigmas papillose c.

O. 5 mm long. Fruit depressed spheroid c. 1.5 mm diam.; pericarp thin, papillose,
loosely adherent. Seed depressed spheroid c. 1.3 mm diam; testa faintly punctuate,
black.

NEW ZEALAND: North Island:— Wellington, 19 Feb. 1950, G. B. Cone (CHR); Te
Kaminara Bay, 21 Nov. 1947, V. D. Zotov (CHR). South Island: — Ben Lomond, 10
Dec. 1943, A. J. Healy(CHR).

Distribution. North and South Islands of New Zealand.

Habitat. ‘Coastal, lowland and montane sandy and rocky places throughout, but
rather local inland’, fide Allan (1961).

Until Paul Aellen’s paper in 1939 Einadia allanii had been confused with E.
triandra (while plants of the latter were generally referred to Rhagodia nutans). The
two species have superficial resemblances but they differ in leaf-shape and in fruit
since the pericarp of E. allanii is thin and dry while that of E. triandra is succulent.
Einadia allanii is evidently closely related to E. trigonos for they are similar in habit
as well as in flower and fruit character. The two are separable on leaf-shape, that of
E. allanii being broadly elliptic (with rounded apex) to orbicular while in E. trigonos
it is bluntly hastate. The situation is complicated by the presence in New Zealand of
the typical subspecies of E. trigonos, of its subspecies stellulata (both presumably
introduced in the past 200 years), and of plants which are intermediate in character
between the typical variants of E. trigonos and E. allanii. It seems likely that these
‘intermediate’ collections are of hybrid origin but this suggestion needs to be sub-
stantiated by field observations since E. trigonos is extremely variable in Australia,
as regards leaf form. This makes its recognition (and of putative hybrids) in New
Zealand uncertain.

Einadia allanii exhibits considerable variability in leaf shape and texture, and in
inflorescence character. I have adopted a rather broad concept of the species since I
am unable to find any clear characters by which the complex could be subdivided.
One variant does stand out and would appear to warrant recognition as a distinct
species, this is a plant found on Little Barrier Island that has an open paniculate in-
florescence the branches of which are destitute of bracts. A plant very similar in form
is also found on the Kermadec Islands. Until further work is done on possible rela-
tives in other islands of the Pacific or in South America it would seem best not to de-
scribe it as a distinct taxon. The following specimens from New Zealand represent
this variant.

Paul G. Wilson, Tribe Chenopodieae

211

LITTLE BARRIER IS.: T. F. Cheeseman (CHR 331646); R. Melville 6572 (CHR
140167).

KERMADEC ISLANDS: Curtis Is., W. R. B. Oliver (CHR 289968); Raoul Is., VL. R.
Sykes 1423/K (CHR 326878).

The variant found on Little Barrier Island probably represents the plant illus-
trated in Cheeseman, Illustrations of the New Zealand Flora vol. 2:t 164 (1914),
under the name Chenopodium triandrum. The variation in morphology was com-
mented on by Allan (1961 p. 228), however, it is not clear what taxa he had in mind
since the names Chenopodium allanii and Rhagodia triandra have been applied
rather imprecisely.

5. Einada hastata (R.Br.) A. J. Scott, Feddes Repert. 89:4 (1978). — Rhagodia
hastata R.Br., Prod. 408(1810). Type: Port Jackson, near Duck River on the
Parramatta Road, R. Brown (holo: BM;? iso: K, MEL, NSW). Figure 4F.

Spreading perennial herb (rarely erect to 1.5 m), branching at base. Branches
quadrangular, when young, with vitreous tubular and vesicular hairs (sometimes pale
yellow) soon glabrous. Leaves opposite or subopposite, somewhat coriaceous and
resinous when dry, glabrous or sparsely pubescent with vitreous vesicular hairs when
young; lamina deltoid to broadly hastate, entire, 10-25 mm long, rounded and often
apiculate at apex, lateral lobes short and obtuse, base cuneate to obtuse; petiole slen-
der, somewhat shorter than lamina. Inflorescence a condensed or slender racemose
panicle with scattered glomerules. Flowers sessile or shortly pedicellate, depressed,
0.5-1 mm diam.; tepals oblong-obovate, somewhat fleshy, thickened and cucullate at
apex, remaining incurved over developing fruit, glabrescent, red and somewhat suc-
culent in fruit; stamens 1-2, glabrous, filaments thickened at base, spreading at an-
thesis; ovary depressed, glabrous, smooth; stigmas sessile, spreading c. 0.5 mm long.
Fruit a berry, depressed, clasped by persistent red succulent tepals, c. 3 mm diam.;
pericarp succulent red. Seed lenticular with rounded margin, 1.5 mm diam., radially
rugulose.

QUEENSLAND: Russell Is., C. T. White 3405 (BRI); Harrisons Pocket, S. T. Blake
3203 (BRI).

NEW SOUTH WALES: Tallebung, G. M. Cunningham 645 (NSW); 8 km S of
Inglewood, L. A. S. Johnson 7216 (NSW).

VICTORIA: Mt Wheeler, A. E. Orchard 2662 (AD); Bendigo, A. C. Beauglehole
42128 (MEL).

Distribution. Victoria, eastern Queensland and New South Wales. Map 25.

Habitat. Shady gullies in rainforest, rocky hill slopes, and open plains.

Einadia hastata may be recognised by its relatively thick, green, almost glabrous,
broadly hastate leaves with rounded apex. The hairs on the young shoots have a vit-
reous appearance unlike the milky colour of those in E. nutans and E. trigonos.
Maiden in Agric. Gaz. New South Wales 8:123-125 (1897) states that the crushed
leaves have an odour of perfectly fresh fish. It varies considerably in habit and in leaf
size and texture but less so in leaf shape and least of all in seed characters.

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Nuytsia Vol. 4, No. 2 (1983)

6. Einadia polygonoides (Murr) Paul G. Wilson, comb. nov.—Chenopodium
triangulare R.Br. var polygonoides Murr, Bull. Herb. Boiss. 4:994 (1904). — C.
triangulate subsp. polygonoides (Murr) Murr, Allg. Bot. Zeitschr. 16:56 (1910). — C.
polygonoides (Murr) Aellen, Mitteil. Ges. Soloth. 8:213 (1928). Type citation: ‘Port
Jackson district NSW, leg. Ernst Betke, 2. 1898, comm. Charles Walter (v. in herb.
Turic.).’ Iso (?): Kogarah (Sydney), Feb. 1898, J. H. Camfield (NSW 143135, ? MEL
88743), see comment below.

C. triangulare R.Br. var. angustifolium Benth., FI. Austral. 5:161 (1870). Type ci-
tation: ‘To this belong the Queensland specimens [of C. triangulare] and some from
New England.’ Lectotype: Armadilla, W. Barton 140 (MEL 80674) lecto. nov.

Spreading weak grey-green herbaceous perennial with numerous stems arising
from woody taproot. Branches slender, striate, very sparsely mealy when young with
vesicular hairs. Leaves opposite or alternate; lamina chartaceous, narrowly oblong-el-
liptic to narrowly hastate, 0.5-1. 5 cm long; apex acute; base obtuse to truncate; upper
surface glabrous or glabrescent; lower surface mealy with white vesicular hairs when
young; petiole slender c. one quarter length of lamina. Inflorescence a compact ter-
minal racemose panicle of dense glomerules of flowers, elongating in fruit to 15 cm
with disjunct glomerules; axis when young with short tubular and globular hairs and
with small gland-tipped hairs. Flowers sessile, 0.5-1 mm diameter; tepals broadly ob-
ovate with rounded apex; stamens 1(2), glabrous, filament thickened at base and
spreading at anthesis; ovary depressed, papillose; style c. 0.1 mm; stigmas slender, di-
varicate, c. 0.5 mm long. Fruit: tepals imbricate, closely enveloping nut, coriaceous,
with circular limb and short narrow claw, glabrous somewhat resinous when fresh,
eventually straw-coloured to black, midrib prominent, pericarp thin, dry, promi-
nently papillose above. Seed lenticular with rounded margin, 1 mm diam.; testa
finely radially rugulose.

QUEENSLAND: Kindon Station, L. S. Smith 514 (BRI); Chinchilla, J. P. Ryan
(BRI).

NEW SOUTH WALES: Marayong, 3 July 1966, R. Coveny (NSW); Narellan, E.
McBarron 13673 (NSW).

SOUTH AUSTRALIA: Arno Bay, Jan. 1920, T. G. B. Osborn (AD); Clapham, 2 Mar.
1962, R. Pearce (ADW).

Distribution. Central-east Queensland to central-east New South Wales; as an
introduction, in Eyre Peninsula and Adelaide, South Australia. Map 28.

Habitat. Appears to favour heavy loamy soils.

Einadia polygonoides may be readily recognised by its narrow leaves, slender
infructescence and free coriaceous tepals with circular limbs. The indumentum of
this species is unusual for Einadia since some of the small multicellular hairs are ter-
minated by a minute spherical cell which appears to be filled with resin.

The type of Chenopodium triangulare var. polygonoides was stated by Murr (1904
l.c.) to have been collected by ‘Betke’ (a misspelling of ‘Betche’). However, a note on
a probable isotype in herb. NSW states that the collector was J. H. Camfield. Murr
(1910) cites the Camfield collection and states that Betche collected it at the same
place and date. Whether there were one or two collections is not clear but it seems
certain that they are of the same population. The type specimen probably represents
a relatively recent introduction to the Port Jackson district.

Paul G. Wilson, Tribe Chenopodieae

213

5. RHAGODIA R.Br.

R.Br., Prod. 408 (1810); A. J. Scott, Feddes Repert. 89:5 (1978). Lectotype: R.
billardieri R.Bt. (= R. baccata (Labill.) Moq.), see Ulbrich (1934 p. 480).

Dioecious shrubs, ± mealy (at least when young) with minute globular or lobed
vesicular hairs, these (in some species) collapsing and becoming scurfy, scaly, or
fusing to form a continuous silvery sheen. Leaves opposite or alternate, simple, entire
or bluntly lobed at base, 2-40 mm long. Flowers small, glomerulate, arranged in pan-
icles or reduced to spike-like thyrses; bracts small and linear or absent; perianth ±
globular c. 1.5 mm diameter, usually mealy outside and glabrous within; tepals 5,
shortly united at base, often spreading and slightly enlarged in fruit, chartaceous or
slightly fleshy. Male flowers: stamens, 5, filaments linear, usually united at base into
a thickened saucer- or cup-shaped disc woolly or puberulous within (rarely glabrous),
anthers exserted; pistillode minute, with or without a pair of minute erect stigmas.
Female flower if terminal to glomerules or branches then with staminodes and a disc
which is woolly within, if lateral then with small usually glabrous staminodes or
these absent; ovary glabrous and globular; stigmas 2, sessile, divaricate, papillose.
Fruit a berry, depressed globular, c. 3 mm diameter, yellow or red; pericarp fleshy
usually with watery interior, viscous on drying. Seed horizontal, depressed globular
or lenticular, 1-1.5 mm diameter; testa crustaceous, black, smooth to rugulose or re-
ticulate; embryo annular; perisperm copious central.

Eleven species, all endemic to Australia (including Tasmania).

Key to species

1 .

2 .

3.

4.

5.

6 .

Leaves small c. 3 mm long; intricately branched spinescent shrubs 2

Leaves more than 3 mm long or plant not spinescent 3

Stamens woolly at base; seed radially verruculose (W.A.) 4. R. acicularis

Stamens glabrous; seed smooth to minutely granular (N.S.W., S.A.,

W.A.) 3. R. ulicina

Perianth glabrous or almost so 4

Perianth mealy or with minute vesicular hairs 5

Leaves linear to narrowly or broadly obovate somewhat

mealy below 5. R. preissii

Leaves narrowly elliptic with minute spherical or saucer-shaped vesicular hairs

(rarely glabrous) 6. R.baccata

Leaves mealy when young, becoming scaly and covered with a silvery sheen;

staminodes present in female flower 6

Leaves mealy or of irregular to spherical vesicular hairs or glabrous (not becom-
ing scaly or forming a sheen) 7

Leaves papery or slightly fleshy, flat 1. R. candolleana

Leaves fusiform or semiterete, fleshy 2. R. crassifolia

7. Indumentum of spherical (or saucer-shaped) vesicular hairs 8

Indumentum of irregularly shaped vesicular hairs 10

8. Leaves oblong to narrowly elliptic, c. 30 mm long, glabrous above, sparsely ves-
icular pubescent below (coastal south-west Australia) 6. R. baccata

Leaves elliptic to deltoid or circular (inland or eastern States) 9

22493 — ( 6 )

214

Nuytsia Vol. 4, No. 2 (1983)

9. Leaves narrow elliptic, lamina mostly 7-10 mm long; female flowers lacking both

disc and staminodes; seed almost smooth (W.A.) 8. R. drummondii

Leaf-shape varied, lamina (3) 5-20 mm long; lateral female flowers with disc and
often staminodes; seed prominently reticulate (central and eastern Australia).

9. R. spinescens

10. Leaves broadly obovate to orbicular or bluntly hastate, flat; inflorescence

pyramidal 11

Leaves linear to elliptic or oblong-elliptic 12

11. Leaves obovate to orbicular, often conduplicate; indumentum of closely packed
glossy, irregularly shaped vesicular hairs (W.A., Shark Bay to

Geraldton) 7. R. latifolia

Leaves broadly ovate to bluntly hastate, flat; indumentum of very minute ves-
icular hairs, dull (S.A., N.T., eastern Australia) 11. R. parabolica

12. Leaves semiterete, somewhat fleshy; panicle narrow, often racemose (W.A.,

Geraldton area) 7. R. latifolia subsp. recta

Leaves elliptic to ovate (inland or coastal south-west) 13

13. Leaves ovate or ovate-oblong, c. 15 mm long, thinly covered below with minute
mealy hairs; flowers in dense slender spikes, sometimes paniculately arranged;
staminodes usually absent in female flower (S.A., W.A., N.T., inland species)

10. R. eremaea

Leaves elliptic, 10-20 mm long, densely covered below with minute vesicular
hairs; flowers in dense panicles (W.A., coastal south-west).

6. R. baccata subsp. dioica

1. Rhagodia candolleana Moq., Chenop. Enum. 10 (1840). — R. baccata var.
candolleana (Moq.) Moq. in DC., Prod. 13/2:50 (1849). Lectotype: Nouvelle hollande,
Labillardiere (holo: P) lecto. nov. (See discussion below.) Figures 1, 2.

Cbenopodium furfuraceum Moq. in DC., Prod. 13/2:64 (1849). Type: Terre de
Diemen — detroit d’ Entrecasteaux, 1802, herb. Delessert (holo: P).

[R. billardieri auct. non R.Br.: Benth., FI Austral. 5:152 (1870) p.p. excluding West-
ern Australian specimens cited.]

[R. baccata auct. pi. non (Labill.) Moq.: e.g. J. Black, FI. S. Austral, ed. 2, 287 (1948);
Beadle et al., FI. Sydney Reg. 192 (1972); J. H. Willis, Handb. FI. Viet. 2:83 (1973).]

[R. billardieri var. congesta auct. non (J. D. Hook.) Benth.: Benth., FI. Austral. 5:153
(1870).]

[R. baccata var. congesta auct. non (J. D. Hook.) A. J. Scott: A. J. Scott, Feddes
Repert. 89:7 (1978).]

Weak spreading shrub or erect and up to 5 m high. Branches striate, mealy when
young. Leaves opposite to alternate, shortly petiolate; lamina chartaceous to cori-
aceous or somewhat fleshy, narrow to broad elliptic to broad ovate or hastate, 10-
25(40) mm long, acute to obtuse; upper surface sparsely mealy to glabrous; lower sur-
face mealy developing a continuous grey to silvery sheen with age or glabrescent. In-
florescence a terminal pyramidal panicle to 15 cm long. Flowers predominantly di-
oecious; perianth fleshy, densely pubescent outside with weak irregularly shaped
hairs, tepals ovate and united near base, midrib thickened. Male flowers: stamens 5,
filaments linear-lanceolate, united at base into a thick saucer-shaped disc shortly
puberulous within; pistillode from minute to normal size with small erect stigmas.
Female flower: staminodes with ovate membranous filaments 1/4 to 1/2 length of

Paul G. Wilson, Tribe Chenopodieae

215

tepals, sometimes terminating in minute antherodes, united at base into a thickened
disc woolly within; ovary glabrous, style arms spreading, c. 0.5 mm long. Berry
clasped at base by the fleshy red perianth. Seed depressed globose, 1.5-2.5 mm diam.,
testa almost smooth to faintly reticulate or rugulose.

Key to subspecies

Leaves elliptic to broadly ovate or hastate, lower surface grey or glabrescent

la. subsp. candolleana

Leaves hastate, acute, lower surface with a silvery sheen lb. subsp. argentea

la. subsp. candolleana

R. baccata var. parvifolia Moq. in DC., Prod. 13/2:50 (1849). Type: Port Jackson,
Durville (holo: P).

Chenopodium furfuraceam Moq. op. cit.

Leaves narrow to broad elliptic, broad ovate, or hastate, obtuse; lower surface
covered with a grey sheen or glabrescent.

NEW SOUTH WALES: 2 mi [3.2 km] N of Bermagui, H. Salasoo 5107 (NSW);
Kogarah, Nov. 1899, J. H. Camfield (NSW).

VICTORIA: Lady Julia Percy Is., A. C. Beauglehole 49662 (MEL); Coringle, N. W.
Wakefield 4274 (MEL).

TASMANIA: Sulphur Creek, Jan. 1954, W. M. Curtis (HO); Coles Bay, 20 Apr. 1930,
L. Rodway (HO).

SOUTH AUSTRALIA: Henley Beach, 26 Mar. 1921, E. H. Ising (AD); Port
Noarlunga, L. D. Williams 5134 (AD).

WESTERN AUSTRALIA: Middle Is., Recherche Archipelago, A. S. Weston 8856
(PERTH); Observatory Is., A. S. Weston 9386 (PERTH).

Distribution. Tasmania, southern New South Wales, Victoria, South Australia,
Recherche Archipelago in Western Australia. Map 29.

Habitat. Coastal situations.

lb. subsp. argentea Paul G. Wilson, subsp. nov.

Frutex omnis argenteo-squamosus. Folia aliquantum hastata 10-25 mm longa,
acuta.

Typus: South Australia; Maralinga Village, 15 July 1972, T. R. N. Lothian 5547
(holo: AD; iso: NCU n.v.).

Shrub closely covered all over with a silvery scaly indumentum. Leaves somewhat
hastate, 10-25 mm long, acute.

SOUTH AUSTRALIA: Ifould Lake, T. R. N. Lothian 5429 (AD); Maralinga, 7 Feb.
1960, H. Turner (AD).

216

Nuytsia Vol. 4, No. 2 (1983)

Distribution. South Australia, North-West Plains.

The subspecies argentea has a distinctive leaf shape and a thick silvery sheen. It is
clearly distinct in its distribution from the typical subspecies and could be separated
(on morphological characters) at the species rank.

Near Maralinga and at Ifould Lake subsp. argentea has an acutely hastate leaf; at
Ooldea has been collected a plant (Spooner 2185, AD) which is similar but has an el-
liptic leaf; also at Ooldea has been collected a plant (J. B. Cleland, 17 Aug. 1939, AD)
with leaves similar to the Spooner specimen but with flowers and indumentum more
like those of Chenopodium desertorum. Until extensive field investigations take
place no satisfactory interpretation of this data is possible.

On the Nullarbor Plain 12 km NW of Cocklebiddy Rhagodia crassifolia may be
found growing with a plant intermediate in morphology between that species and R.
candolleana subsp. argentea. This putative hybrid (P. G. Wilson, 11829, PERTH)
suggests that R. candolleana subsp. argentea is also found in that district.

Lectotypification of the name Rh^odia candolleana. The correct application of the
name Rhagodia baccata, and the reasons for rejecting it for the species occurring in
eastern Australia, are discussed under that name. The lectotypification of the
alternative name, R. candolleana, also requires an explanation since it was based on
mixed material. Moquin (1840) cited the type of R. candolleana as “In nova Hol-
landia (v.s. in herb. Mus. Paris. DC., et Vindob.)”. In 1849, under R. baccata var.
candolleana (Moq.) Moq., he wrote “v.s. in h. Labil. et Mus. Paris.” I have seen four
sheets which appear to be syntypes of this species, their labels are as follows:

A. Herb. P

Label (1) Rhagodia candolleana Moq./Nouvelle hollande/Labiiiardfere/Herb.
Poiret.

(2) Rhagodia Candolleana Moq./Nova hollandia/herb. Vindob.

(3) Herb. Mus. Paris/Herbier Moquin-Tandon/(Monographies du
Prodrome)

B. Herb. P

Label (1) Rhagodia/Nova Holland/ Drummond No. 209.

(2) Rhagodia hastata R.Br./(non)/Nova hollandia/herb. Vindob.

(3) Herbarium Moquinianum/Rhagodia baccata Moq./var.
Candolleana/ (R. candolleana Moq.) /nova Holland. (La Bill).

(4) Herb. Mus. Paris/Herbier Moquin-Tandon/(Monographies du
Prodrome)

C. Herb. FI

Rhagodia Candolleana Moq. [In Moquin’s handwriting]

D. Herb. G-DC

Label (1) Nouv Hollande cote d’orientale/Museum de Paris 1821

(2) baccata/Rhagodia Billardieri R.Br./var.CandoWeana/Moq.

(3) Rhagodia/Nouv Hollande cote Merid./Mus de Paris 1821

Paul G. Wilson, Tribe Chenopodieae

217

holo IVPE

KEW NEGATIVE

NO, 171 90
' mj

frt'l »*'! o-ll

■’■Jt) 2U 34 4-U

‘L^

oflt n#

ii'* 0^* ■y¥ 01 02

»0 I2» !3'0

01

14.0

Figure 1. Lectotype of Rhagodia candolleana Moq. in herb. P.

218

Nuytsia Vol. 4, No. 2 (1983)

Sheets A and D contain only specimens of the eastern Australian plant here
referred to Rhagodia candoUeana. Sheets B and C each contain a small vegetative
specimen of R. candoUeana and a larger specimen of Tetragonia implexicoma (Miq.)
J. D. Hook. (Aizoaceae). I have selected sheet A in herb. P. (Figure 1) as the
lectotype since it bears Moquin’s handwritten determination and is of unmixed ma-
terial. It is certainly one of the specimens studied by Moquin when describing the
species. The sheet B in herb. P has, as noted above, a Drummond label no. 209. This
is misplaced since James Drummond could not have collected either of the two speci-
mens on the sheet; the Drummond no. 209 collection in herb. MEL is of Rhagodia
baccata sensu stricto (i.e. the Western Australian plant) from which the Drummond
label in herb. P had evidently strayed.

Rhagodia candoUeana subsp. candoUeana varies considerably in leaf form, texture
and sculpturing of the seed (faintly rugose in Tasmania to faintly reticulate in South
Australia); it also varies in stature, being a shrub to 5 m high in Tasmania while
rarely exceeding 1 m on the Australian mainland. However, there are insufficient
supporting characters with which to establish further infraspecific taxa. In South
Australia it is often not possible to clearly distinguish R. candoUeana from the
coastal variant of R. crassifolia, however, since elsewhere these two species appear to
be clearly separable I have continued to recognise them.

Although predominantly coastal in its distribution subsp. candoUeana has been
collected at Mt Gambier in South Australia. J. M. Black (1948) recorded it inland
along the R. Murray but the relevant specimens are all of the inland variant of R.
crassifolia. Black also recorded Rhagodia baccata var. linearis for South Australia,
however, material so determined in his herbarium is either Suaeda australis (R.Br.)
Moq. or R. crassifolia.

Rhagodia candoUeana is nearly always dioecious but populations of apparently
hermaphrodite plants do occur. Male plants sometimes have prominent pistillodes
and these frequently enlarge to form berry-like structures which are probably galls.

The peculiar indumentum of Rhagodia candoUeana (and of R. crassifolia) is made
up of extremely thin-walled mealy hairs of irregular shape. These soon collapse and
fuse together to form a metallic sheen similar to that found in some species of
Atrip lex.

The name Chenopodium congestum J. D. Hook, is based on a specimen of C. mu-
rale L., q.v. However, as a variety of Rhagodia billardieri or of R. baccata it has been
misapplied by Bentham (1870) and Scott (1978a) to variants of R. candoUeana in
which the leaves tend to be rhombic or hastate and the inflorescence somewhat con-
gested.

The type of Chenopodium furfuraceum is a male specimen of Rhagodia
candoUeana. It was not seen by Bentham (1870) who thought it might be conspecific
with C. auricomum.

2. R. crassifolia R.Br., Prod. 408 (1810). Lectotype: Goose Island Bay, larger
island, 15-16 Jan. 1802 R Brown, dist. no. 3038 (holo: BM) lecto. nov.

Kparvifolia Moq. in DC., Prod. 13/2:52 (1849). Type: Ad Cygnorum fluvium, 1843, J.
Drummond 715 (holo: P).

Paul G. Wilson, Tribe Chenopodieae

219

Shrub to 1.5 m high, divaricately branched. Leaves mostly alternate, fleshy, linear
to narrow-elliptic in outline, fusiform or semiterete, often conduplicate or channelled
above, falcate when thick, 1-2(4) cm long, mealy when young, becoming glabrescent
above, hairs collapsing and fusing to form a metallic sheen. Inflorescence pyramidal
to 5 cm long or reduced to a short spike. Flowers and fruit similar to that in
R. candolleana. Seed depressed globose, 1.5 mm diam., very faintly reticulate or
rugulose.

VICTORIA: 16 mi [25.7 km] NNW of Underbool, A. C. Beauglehole 40470 (MEL); 5
mi [8 km] E of Murrayville, 29 Aug. 1955, J. H. Willis (MEL).

SOUTH AUSTRALIA: Nundroo, A. G. Spooner 2194 (AD); Port Lincoln, 17 Dec.
1941, J. B. Cleland (AD).

WESTERN AUSTRALIA: Corbett Is., Recherche Archipelago, R. D. Royce 6189
(PERTH); 12 km E. of Mundrabilla, P. G. Wilson 11832 (PERTH).

Distribution. North western Victoria westwards to near Albany on the south coast of
Western Australia. Map 30.

Habitat. Principally a coastal species but also inland where usually on calcareous
soil; often in a mallee community.

Rhagodia crassifolia intergrades with R. candolleana in coastal regions of South
Australia and many collections cannot be confidently placed in either species. Near
Cocklebiddy in the southern Nullarbor Plain of Western Australia R. crassifolia has
been found growing with an apparent hybrid (P. G. Wilson 11834, PERTH) between
that species and R. candolleana subsp. argentea.

The coastal variant of R. crassifolia differs somewhat from that found in the
Murray-Mallee region of South Australia and adjacent areas of Victoria; this inland
variant has a strongly divaricate habit and longer more slender leaves. It was princi-
pally this variant that J. M. Black (1948) incorrectly referred to under the name
R. baccata var. linearis.

Moquin (1849 loc. cit.) based his concept of R. crassifolia largely on a specimen
collected by Alan Cunningham on Dirk Hartog Island. This collection is, in fact, of a
distinct species, R. latifolia (Benth.) Paul G. Wilson. The specimen of R. crassifolia
collected by James Drummond in Western Australia, being considered by Moquin to
be specifically distinct from that of Cunningham, was described as a separate
species, R. parvifolia. Although Moquin cited its locality as ‘Ad Cygnorum fluvium’,
i.e. from Swan River Western Australia, the specimen was probably collected near
the Fitzgerald River mouth since it closely matches other specimens from that area
while the species is not found on the west coast of Western Australia.

Bentham (1870) and Ewart and Davies (1917) recorded R. crassifolia from North-
ern Australia, in both cases citing a specimen from Sturt Creek collected by F.
Mueller. This specimen is probably a syntype of Chenopodium nitrariaceum (F.
Muell.) Benth. but is not conspecific with the lectotype of that species; it belongs in
fact to R. eremaea.

3. Rhagodia ulicina (Gand.) Paul G. Wilson, Nuytsia 4:53 (1982.)— Chenopodium
ulicinum Gand., Bull. Soc. Bot. France 66:224(1919). Type: Mt Lyndhurst, April
1900, M. Koch 366 (holo: LY; iso : PERTH). Figure 5F.

220

Nuytsia Vol. 4, No. 2 (1983)

Dense intricately branched shrub 0.5-1 m high, dioecious. Branches divaricate,
spinescent, mealy when young with close dense covering of small branched vesicular
hairs. Leaves alternate, often clustered, fleshy, elliptic, 2-5 mm long, closely vesicular
mealy, narrowed at base into an indefinite petiole. Inflorescence a few-flowered nar-
row condensed cyme (sometimes spinescent) with 1-4 lateral and a large terminal
flower, or reduced to a single flower. Perianth depressed globose 1.5 mm diam.,
mealy outside with small branched vesicular hairs; tepals broadly ovate, united at
base, eventually spreading beneath fruit. Male flower: stamens 5, filaments slender,
glabrous, disc inconspicuous, anthers shortly exserted c. 0.5 mm long; pistillode min-
ute. Female flower: staminodes represented by 5 scarcely observable antherodes c.
0.5 mm long; ovary glabrous; style absent to short and c. 0.5 mm long, glabrous or
sparsely hirtellous; stigmas spreading, slender subulate, pilosulose. Fruit baccate, at
first enveloped by perianth; pericarp succulent, appearing inflated on drying, free
from seed. Seed lenticular with rounded margin, c. 1.5 mm diam., testa smooth to
minutely granular, black.

NEW SOUTH WALES: Fowlers Gap, S. Jacobs 2268 (NSW); 20 mi [32.2 km] N of
Balranald, W. E. Mulham W518 (NSW).

VICTORIA: Yatpool area, J. H. Browne 117 (PERTH).

SOUTH AUSTRALIA: Mt Grainger, R. J. Chinnock 1479 (AD); Wilkawillina Gorge,
T. R. N. Lothian 5126 (AD).

WESTERN AUSTRALIA: 19 km NNW of Mt Glass, K. Newbey 5609 (PERTH); 12
km NW of Cocklebiddy, P. G. Wilson 11827 (PERTH).

Distribution. Eastern New South Wales and north-eastern Victoria to Northern Eyre
Peninsula in South Australia westwards to the south-east of Western Australia
around the Bremer Range. Map 31.

Habitat. Generally rocky situations often over limestone, but has been recorded from
clay flats.

This species has been confused in herbaria with both R. spinescens and Cheno-
podium nitrariaceum. It was included by Bentham (1870) under the former name
(i.e. the specimens cited as ‘Murray river, F. Mueller’, ‘Gawler Ranges, Sullivan’, and
‘Barrier Range, Victorian Expedition’), however, his description of R. spinescens was
based not only on that species and R. ulicina but also on Chenopodium desertorum.
Gandoger (1919 loc. cit.) recognised R. ulicina as being a distinct species but his de-
scription was very brief and suggested that he was unaware that the plant was di-
oecious, shrubby, and had baccate fruits, for these characters establish the species as
a member of the genus Rhagodia and not Chenopodium. Scott (1978b) included R.
ulicina (as Chenopodium) with C. nitrariaceum under Chenopodium sect.
Rhagodioides Benth.; however, Scott’s sectional description is inapplicable to R.
ulicina and the two species are not closely related.

Rhagodia ulicina may be distinguished from R. spinescens and C. nitrariaceum by
its intricately branched habit and usually by its small fleshy leaves. Some specimens
of C. nitrariaceum resemble R. ulicina in leaf shape but differ in indumentum: that
of C. nitrariaceum is scurfy and forms a silvery sheen with age, while the vesicular
hairs of R. ulicina are irregularly lobed and remain distinct.

Rhagodia ulicina is most similar to R. acicularis (an endemic of the Wongan Hills,
Western Australia); the differences between them are give under the latter species.

Paul G. Wilson, Tribe Chenopodieae

221

4. Rhagodia acicularis Paul G. Wilson, Nuytsia 4:51 (1982). Type: Monk’s Well
Gully, Wongan Hills, 15 Feb. 1980, P. G. Wilson 11713 (holo: PERTH; iso: CANB).
Figure 5E.

Compact intricately branched rounded shrub c. 0.5 m high, closely covered with
minute vesicular hairs which retain their shape with age, dioecious. Branchlets
spinescent. Leaves alternate, small, somewhat fleshy, narrowly elliptic, 2-5 mm long,
glabrescent above, closely vesicular puberulous below. Inflorescence spicate (or
shortly paniculate) with a spinescent axis c. 10 mm long; flowers few, sessile, sub-
tended by a minute bract. Male flowers broadly turbinate, c. 1 mm long; tepals 5,
divided to base, narrowly obovate, incurved, closely vesicular hairy outside, glabrous
within; stamens 5; filaments slender-terete, lanate towards base, united into a thick
cup-shaped disc; anthers shortly exserted, c. 0.5 mm long; pistillode minute. Female
flowers spheroidal, c. 1.2 mm diam.; tepals narrowly obovate not enlarging in fruit;
staminodes absent; ovary glabrous; style absent; stigmas slender, papillose, c. 0.3 mm
long. Terminal flower on female inflorescence sterile, densely filled with woolly hairs
arising from staminodes; pistillode minute with 2-3 erect glabrous stigmas. Fruit bac-
cate, depressed spherical, clasped around margin by tepals but exposed at apex and
towards base in inter-tepalar spaces; pericarp succulent, adherent to seed. Seed len-
ticular with rounded margin, 1.5 mm diam., radially verruculose.

WESTERN AUSTRALIA: Wongan Hills, K. F. Kenneally 2384 and 6448 (PERTH);
ibid., P. G. Wilson 11712 (PERTH).

Distribution. Wongan Hills, south-western Western Australia.

Habitat. Red soil on gravelly lateritic slope in eucalypt woodland.

Rhagodia acicularis closely resembles R. ulicina but differs in the following charac-
ters: 1) Terminal flower of female inflorescence sterile (not large and fertile), 2) Seed
radially verruculose (not smooth or minutely granular), 3) Male flower with woolly
stamens united into a cup-shaped disc (not glabrous and with an inconspicuous disc).
It is a typical member of the genus and evidently closely related to R. spinescens R.
Br. and R. drummondii Moq.

5. Rhagodia preissii Moq. in DC., Prod. 13/2:49 (1849).— Chenopodium preissii
(Moq.) Diels, Bot. Jahrb. 35:181(1904). Type: Near York, Western Australia, Preiss
1252 (iso: K, MEL).

R. obovata Moq., Chen. Mon. Enum. 10(1840). Type: Nouv. Holland, cote meridi-
onale, 1821 (holo: P).

[R. linifolia auct. non R.Br.: Nees in Lehm., PI. Preiss. 1:633(1845).]

Shrub to 2 m high. Branches slender, when young costate and sparsely mealy.
Leaves chartaceous (slightly fleshy when fresh), to coriaceous (subsp. obovata), lin-
ear to broadly obovate or spathulate, 3-4 cm long, gradually tapering at the base into
a slender petiole, apex rounded to acute, lower surface sparsely and minutely mealy
when young. Inflorescence a pyramidal panicle c. 5 cm long, branches slender,
flowers in disjunct glomerules. Flowers spherical, c. 1 mm diam; tepals almost orbicu-
lar, glabrous and eciliate (or sparsely mealy in subsp. obovata) enlarging slightly in
fruit. Male flowers: filaments united into a saucer-shaped disc, somewhat pilose at
base; pistillode minute (c. 0.5 mm high) with a pair of short erect subulate stigmas.
Female flowers: terminal flowers usually sterile with prominent linear staminodes

222

Nuytsia Vol. 4, No. 2 (1983)

densely pilose at base; pistillode ovoid, 0.3-1 mm high: non-terminal flowers fertile;
staminodes absent; ovary with a pair of subulate stigmas. Berry depressed spherical,
c. 3 mm diam. Seed c. 1.5 diam.; testa glossy with faint radial ribbing.

Key to subspecies

Leaves linear to narrow obovate, usually chartaceous 5a. subsp. preissii

Leaves broadly obovate, coriaceous 5b. subsp. obovata

5a. subsp.preissii

Leaves linear to narrow obovate or spatbulate chartaceous (to coriaceous), flat or
cymbiform. Perianth glabrous.

SOUTH AUSTRALIA: Barton, N.T. Burbidge 1866 (CANB); Dublin, J. C. Noble 84
(AD).

WESTERN AUSTRALIA: Merredin, M. Koch 2882 (NSW); 32 km NNE of Stokes
Inlet, A. E. Orchard 1645 (CANB).

Distribution. West coastal and south-west Western Australia, southern South Aus-
tralia. Map 32.

Habitat. Often in deep sand.

5b. subsp. obovata (Moq.) Paul G. Wilson, comb, et stat. nov. — R. obovata Moq.,
Chen. Mon. Enum. 10 (1840). Type: Nouv. holland. cote meridionale, 1821 (holo: P).

Leaves broadly obovate, gradually to abruptly narrowed at base into a slender
petiole c. 1/3 length of leaf, in all 1.5-3 cm long, coriaceous, glabrescent to thinly
mealy, apex rounded to truncate, apiculate. Perianth glabrous to sparsely mealy.

WESTERN AUSTRALIA: Dorre Is., A.S. Weston 10496 (PERTH); Kalbarri to Red
Bluff, R.J. Hnatiuk 760494 (PERTH).

Distribution. Shark Bay-Geraldton region of Western Australia. Map 32.

Habitat. Frequently growing over limestone.

The application of the names Rhagodia dioica and R. obovata is difficult to deter-
mine due to intergradation of the typical variants of these species with R. baccata
and R. preissii. The type of R. obovata comes from Dirk Hartog Island where the
plant has leathery, broadly obovate leaves and glabrous flowers. On the mainland
around Shark Bay the leaves are somewhat narrower and the flowers slightly mealy,
while further south the mealiness increases and the leaf shape varies from obovate
(near Geraldton) to elliptic (near Perth). Near Perth the flowers and leaves are
densely mealy, and here the plant intergrades with R. baccata; this variant of
intermediate aspect has been described as the species R. dioica. In addition to this
apparent intergradation, R. obovata also passes, to the north and east of Shark Bay,
into typical R. preissii with thin narrow leaves and glabrous flowers. I have, in this
paper, recognised R. baccata as a distinct species because of its normally clearly dis-
tinguishable morphological characters while R. obovata I am including as a

Paul G. Wilson, Tribe Chenopodieae

223

subspecies of R. preissii whose typical subspecies is also clear cut in its characters.
Since the syntypes of R. dioica come near to the typical variant of R. baccata I am
treating them as conspecific.

6. Rhagodia baccata (Labill.) Moq. in DC., Prod. 13/2:50 (1849). — Chenopodium
baccatum Labill., Nov. Holl. PI. Sp. 1: 71, t. 96 (1805). — R. billardieri R. Br., Prod.
408 (1810), nom. illeg. Type “Habitat in capite Van-Diemen.” Lectotype: FI (see
notes below) lecto. nov. Figures 2, 3.

Rhagodia dioica Nees in Lehm., PI. Preiss. 1:636 (1845). Lectotype: Fremantle,
Preiss 1253 (holo: K; iso: MEL) lecto. nov.

Rhagodia radiata Nees in Lehm., PL Preiss. 1:637 (1845). — R. billardieri var. linearis
Benth., FI. Austral. 1:153 (1870). — R. baccata var. linearis (Benth.) J. Black, FI. S.
Austral. 683 (1929); Ewart, FI. Viet. 449 (1931). Type citation: ‘In elevatis arenosis
juxta urhiculam Freemantle, Decembri a 1838. Herb. Preiss. No. 1250’. Isotype: MEL
80680, 80681, 80682.

Rhagodia cygnorum Gandoger, Bull. Soc. Bot. France 66:224 (1919). Type:
Fremantle, 15 March 1899, A. Morrison (holo: LY).

Spreading shrub to 2 m high. Branches slender, costate, mealy when young.
Leaves opposite or alternate shortly petiolate; lamina chartaceous (somewhat fleshy
in subsp. dioica), narrowly oblong-elliptic to elliptic, 1-4 cm long, 3-8 mm wide, apex
rounded to obtuse and minutely apiculate, margin often recurved, upper surface
smooth and glabrous or sparsely mealy, lower surface with prominent midvein,
sparsely to densely mealy with saucer-shaped collapsed vesicular hairs, rarely
glabrous (minutely mealy in subsp. dioica). Inflorescence an open terminal panicle;
flowers clustered in sessile lateral dichasial glomerules. Flowers predominantly di-
oecious, globular, 1.5 mm diam.; perianth densely covered with irregularly shaped
vesicular hairs (rarely glabrous); lobes broadly ovate to orbicular, fleshy in centre.
Female plant: terminal flower of main and lateral branches of panicle with 5 mem-
branous lanceolate staminodes 1 mm long united into a fleshy saucer-shaped disc
densely lanate within, pistil (or pistillode) glabrous with a pair of short erect styles;
lateral flowers (which predominate) glabrous within, lacking staminodes and disc,
ovary globular, glabrous with a pair of short spreading subulate styles. Male plant:
staminal filaments narrowly lanceolate to linear, glabrous; disc saucer-shaped, fleshy;
pistillode ellipsoidal, minute (c. 0.5 mm high) with a pair of short stylar processes.
Fruiting perianth spreading to retlexed with somewhat hardened midrib, pale fawn
in colour, persistent; berry depressed globular, red, pericarp fleshy. Seed lenticular,
c. 1.8 mm diam.; testa glossy, minutely and irregularly tuberculate.

Key to subspecies

Leaves chartaceous with recurved margin (on drying), mealy beneath with discrete
saucer-shaped hairs 6a. subsp. baccata

Leaves fleshy, margin not recurved, mealy beneath with very small hairs

6b. subsp. dioica

6a. subsp. baccata

Rhagodia radiata Nees in Lehm., op. cit. (1845).

224

Nuytsia Vol. 4, No. 2 (1983)

Rhagodia baccata var. angustifolia Moq. in DC., Prod. 13/2:50 (1849). Type citation:
‘v.s. in h. Poir. Mus. Vindob. et DC., comm, a cl. Labill. Cunningh. et F. Bauer.’
Lectotype: Nova Hollandia. Ex herb. Labill. (P; iso: FI) lecto. nov. (see notes below).

R. cygnorum Gandoger, op. cit. (1919).

Leaves narrowly oblong-elliptic to oblong-elliptic, 2-4 cm long, 4-8 mm wide, char-
taceous, margin recurved, lower surface sparsely to densely mealy with saucer-
shaped collapsed vesicular hairs, rarely glabrous.

WESTERN AUSTRALIA: Nornalup Inlet, 15 Jan. 1947, M. B. Johnson (PERTH);
Recherche Archipelago, R. D. Royce 6280 (PERTH); 31 mi [49.9 km] S of Perth, R.
A. Saffrey 877 (PERTH).

Distribution. Western Australia from Guilderton (80 km N of Perth) south and east-
wards around the coast to Cape Arid. Map 31.

Habitat. Coastal or near coastal situations.

This subspecies (under various names) has frequently been considered to be
conspecific with Rhagodia candolleana. It may be readily distinguished by its thin,
narrow leaves with recurved margins and by the absence of staminodes in the female
flowers (when lateral in position). A further distinction lies in the indumentum of
the leaves which consists of collapsed vesicular hairs circular in outline. In R.
candolleana the hairs are irregular in shape, very thin walled, and when collapsed
fuse to form a silvery sheen. The species are both found in the Recherche Archipel-
ago where they retain their distinctive characters; their distribution is otherwise
quite distinct.

Almost glabrous plants of R. baccata subsp. baccata are found at some localities
but this variant grades into the typical one with a mealy indumentum and there is no
reason to segregate it as a distinct taxon.

Lectotypification of Chenopodium baccatum. Labillardiere (1805 loc. cit.) gave as
the provenance of Chenopodium baccatum ‘Habitat in capite Van-Diemen’, i.e. from
the south coast of Tasmania. Material collected by Labillardiere in herb. FI, and so
determined by him, is mounted on four sheets; these contain some specimens which
correspond to the Tasmanian plant and others which could only have been collected
on the south-west coast of Western Australia. The plants from the two areas I con-
sider to be specifically distinct (they have been generally considered by earlier
workers to be distinct at either the level of species or variety); therefore it is import-
ant that C. baccatum be lectotypified.

Three of the four sheets mentioned above were re-identified by Moquin, presum-
ably prior to 1849 when he cited the specimens in his treatment of the
Chenopodiaceae in De Candolle’s Prodromus vol. 13/2. The sheet containing the
largest number of specimens was determined by Moquin as R. baccata, a second
sheet as R. baccata var. candolleana, and a third as R. baccata var. angustifolia.
Moquin had therefore selected one of the four sheets as being representative of the
typical variety.

Labillardiere refers to the leaves as being ‘lanceolatis’ and the inflorescence as
‘racemis compositis’; the fruit was also described. The specimens which match the
Western Australian species have narrowly elliptic leaves and bear large inflor-
escences; they correspond well with Labillardi§re’s description and are a good match

Paul G. Wilson, Tribe Chenopodieae

with his illustration (tab. 96). The specimens which match the Tasmanian species
have short elliptic to obovate leaves and are either vegetative or bear short racernes
of young flowers; they do not agree with either Labillardiere’s illustration or descrip-
tion.

The lectotype of Chenopodium baccatum is, therefore, to be chosen from among
those specimens collected by Labillardiere in Western Australia and preferably from
those selected by Moquin as being typical of the species. The sheet determined as
Rbagodia baccaia by Moquin contains Labillardiere’s manuscript description, four
pieces of the Tasmanian plant, none of which corresponds to the illustration, and
two pieces of the Western Australian, one of which is in flower and the other in fruit
(Figure 2). As this fruiting specimen is an excellent match with tab. 96 (it was prob-
ably the one illustrated) and with the description, I am selecting it as the lectotype
of Chenopodium baccatum. The Tasmanian material I am referring to Rbagodia
candolleana Moq. (q.v.) a species which also requires lectotypification. I have seeri a
photograph of a specimen collected by Labillardiere in herb. PH named by him
Chenopodium baccatum. This is of the Western Australian plant and therefore of
Rbagodia baccata sensu lectotypica.

Lectotypification of Rbagodia baccata var. angustifolia. In the Decandolle Her-
barium at Geneva (G— DC) there are two specimens which appear to be syntypes of
this variety. They are as follows:

(1) Lewis-Land, Feid. Bauer

(2) King George Sound, A. Cunningham

In the Paris Herbarium (P) there is one specimen ‘Ex herb. LabilL’ which came
from Moquin’s herbarium and which was seen by him during the writing of the
Chenopodiaceae for Decandolle’s Prodromus vol. 13/2.

In the Florence Herbarium (FI) there is a specimen collected by Labillardiere
which has been determined by Moquin as Rbagodia baccata var. angustifolia.

Each of the four collections cited above is of the variant of R. baccata found on the
south coast of Western Australia between the Recherche Archipelago and Albany. As
lectotype of var. angustifolia I have selected the Labillardiere specimen in herb. P,
(Figure 3) of which the specimen in herb FI is probably a duplicate.

6b. subsp. dioica (Nees) Paul G. Wilson, comb, et stat. nov.—R. dioica Nees in
Lehm., PI. Preiss. 1:636 (1845). Lectotype: see above.

Leaves elliptic, 10 x 3 - 20 x 7 mm, obtuse, fleshy, coriaceous on drying, upper sur-
face glabrescent, lower surface densely white mealy with very small hairs.

WESTERN AUSTRALIA: Garden Island, May 1946, G. R. Meadly (PERTH);
Fremantle, P. G. Wilson 8693 (PERTH).

Distribution. Western Australia, Geographe Bay northwards to Fremantle. Map 30.

Habitat. Coastal.

This subspecies is sympatric over part of its range with the typical subspecies and
plants intermediate in form between the two taxa may be found. It has thicker leaves

226

Nuytsia Vol. 4, No. 2 (1983)

Figure 2. Lectotype of Rhagodia baccata (Labill.) Moq. Mixed collection made by Labillardiere of R.
baccata and R. candolleana. The left-hand, the upper central, and the lower specimen in the right-
hand cluster are of R. candolleana. The lower central specimen and the upper specimen of the right-
hand cluster are of R. baccata. The lectotype of R. baccata is indicated by an arrow. Photo of sheet in
herb. FI.

Paul G. Wilson, Tribe Chenopodieae

227

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Figure 3. Lectotype of Rhagodia baccata var. angustifolia Moq. in herb. P.

228

Nuytsia Vol. 4, No. 2 (1983)

than subsp. bsccata and a more dense indumentum which is made up of smaller and
less well-defined bladder hairs. The type is of a variant of subsp. dioica rather close
morphologically to the typical subspecies; that is the leaves are thinner and lep
furfuraceous. At the other extreme are found plants with very fleshy leaves similar in
shape and size to those found in R. crassifolia.

7. Rhagodia latifolia (Benth.) Paul G. Wilson, comb, et stat. nov.— R. crassifolia
var. latifolia Benth., FI. Austral. 5:155 (1870). Type: Dirk Hartog Is., Jan 1822, A.
Cunningham 321 (holo: K; iso: MEL).

Shrub to 1.5 m high, dioecious. Branches firm, densely covered when young with
compact spherical or irregularly shaped vesicular hairs. Leaves opposite or alternate,
densely covered all over with compact vesicular hairs; lamina coriaceous to fleshy,
linear to broad elliptic or orbicular, c. 1 cm long; petiole slender c. 5 mm long. Inflor-
escence terminal, pyramidal or slender, glomerules scattered along its branches.
Flowers depressed globular c. 1.5 mm diam; tepals obovate, cucullate, densely vesicu-
lar outside, not enlarging in fruit. Male flower: filaments united into a cup-shaped
disc woolly within; pistillode minute c. 0.2 mm long without stigmas. Female flower
when terminal in position with short membranous staminodes and densely woolly
disc, when lateral in position without staminodes and glabrous within; stigmas
divaricate. Berry depressed globular c. 3.5 mm diam., exceeding the spreading peri-
anth which is pale red within. Seed lenticular, 1.3 mm diam., with faint radial
rugulosity.

Key to subspecies

Leaves elliptic to orbicular; inflorescence pyramidal 7a. subsp. latifolia

Leaves semiterete; inflorescence slender 7b. subsp. recta

7a. subsp. latifolia

Divaricately branched shrub. Leaves coriaceous, elliptic to orbicular, often con-
duplicate and then lunate to semicircular, densely covered all over with compact
spherical vesicular hairs. Inflorescence pyramidal.

WESTERN AUSTRALIA: Dirk Hartog Is., A. S. George 11580 (PERTH); Dorre Is.,
A. S. Weston 10482 (PERTH).

Distribution. Shark Bay south to Murchison River. Map 29.

Habitat. Sand dunes, frequently associated with limestone.

7b. subsp. recta Paul G. Wilson, subsp. nov.

Rami ascendentes, graciles. Folia crassiuscula, semiteretia, c. 10 mm longa, 1 mm
lata, supra plana glabra, infra dense farinosa cum pilis minutis vesicularibus,
irregulariter formatis. Inflorescentia in paniculam gracilem spiciformen disposita.

Typus: Victoria District, Geraldton, Jan 1901, Diels et Pritzel 212 (holo: PERTH).

Branches ascending, slender. Leaves fleshy, semiterete, c. 10 mm long, 1 mm wide,
± flat and glabrous above, densely mealy below with very small irregularly shaped
vesicular hairs. Inflorescence a slender, spike-like panicle.

Paul G. Wilson, Tribe Chenopodieae

229

WESTERN AUSTRALIA: Mt Hill, Oldfield (MEL 87904); Geraldton, C. H.
Ostenfeld343 (PERTH).

Distribution. Western Australia; near Geraldton north to Murchison River. Map 29.
Habitat. Sand dunes.

The typical variant of Rhagodia latifolia is very distinct in appearance since the
orbicular leaves become conduplicate (at least on drying) and are then. lunate in
shape. It evidently grades to the south into subsp. recta (in which the leaves are
semiterete), since plants intermediate in form have been collected.

Rhagodia latifolia is quite distinct from R. crassifolia, under which name it was
placed by Bentham (1870), for in the latter species the hairs of the leaves soon col-
lapse and coalesce to form a silvery sheen while the female flowers possess prominent
staminodes. Material of subsp. recta was considered by Bentham to represent typical
R. crassifolia, a view evidently based solely on the leaf shape.

The infraspecific epithet is taken from the Latin word rectus, straight, referring to
the straight branches of this plant.

8. Rhagodia drummondii Moq. in DC., Prod. 13/2:52 (1849). Type: Drummond
133 (P).

Intricately branched, loose, dioecious shrub up to 1.5 m high. Leaves opposite or
alternate, somewhat coriaceous (slightly fleshy when fresh), narrowly elliptic to ellip-
tic, shortly petiolate, 7-10(15) mm long, 2-3(4) mm wide, apex acute, base cuneate,
upper surface glabrescent, lower surface densely covered with pale grey silvery
spherical to saucer-shaped vesicular hairs. Influorescence a short terminal panicle to
2.5 cm long, frequently reduced to a short spike-like thyrse. Flowers sessile, de-
pressed globular to broadly turbinate, solitary or in small glomerulate dischasia;
tepals broadly ovate, shortly united at base, bluntly keeled towards apex, densely
covered with vesicular hairs. Male flowers: staminal filaments united into a fleshy
disc which is woolly within, pistillode minute without trace of stigmas. Female
flowers: terminal flowers with minute staminodes and disc, slightly woolly within;
lateral Rowers glabrous within and without staminodes or disc, stigmas slender.
Fruiting perianth at first clasping fruit hut eventually spreading, c. 4 mm diameter,
pale fawn or pale red within; berry depressed globular; seed lenticular, c. 1.8 mm
diam., faintly radially rugulose, glossy.

SOUTH AUSTRALIA: 12 km E of the Serpentine Lakes, 28°30'S, 129°04'E, D. E.
Symon 12540 (ADW); 9 km N of Connie Sue Crossing on S.A.-W.A. border, 28°25'S,
129°00'E, D. E. Symon 12621 (ADW).

WESTERN AUSTRALIA: Dowerin, A. M. Ashby 2849 (AD); 14.7 mi [23.7 km] N. of
Winchester, M. D. Tindale 1289 (NSW).

Distribution. Inland southern Western Australia eastwards to the extreme west of
South Australia. Map 33.

Habitat. Somewhat saline inland areas, also in Eucalyptus salmonophloia (Salmon
Gum) Woodland.

22493 — ( 7 )

230

Nuytsia Vol. 4, No. 2 (1983)

Rhagodia drummondii is similar to R. spinescens but differs most obviously in the
leaf shape (always elliptic) and in the absence of staminodes in the lateral female
flowers. It was considered by Bentham (1870) and by Scott (1978b) to be a form of
R. crassifolia, however, the two are not closely related; they differ very greatly in in-
dumentum (the hairs of R. crassifolia collapsing to form a continuous metallic sheen)
and in flower morphology, particularly in the presence of prominent staminodes in
the female flowers of R. crassifolia.

9. Rhagodia spinescens R.Br., Prod. 408 (1810). Type: Inlet XIV [St. Vincent
Gulf], R. Brown (holo: BM; iso: K). Figure 5D.

R. spinescens var. deltophylla F. Muell., Rep. PI. Babbage’s Exped. 19 (1859).— R.
deltophylla (F. Muell.) A. J. Scott, Feddes Repert. 89:10 (1978). Type citation: Lake
Campbell. Lectotype: N.E. of Lake Gairdner, Hersch [Herschel] Babbage.’ (MEL
87960) lecto. nov. (see note below).

Intricately branched shrub to 1-5 m high. Branches divaricate, frequently
spinescent, densely mealy when young. Leaves opposite or subopposite, chartaceous
to coriaceous, mealy all over with discrete spherical or saucer-shaped hairs, lamina
ovate to deltoid or hastiform 0.5-2 cm long, apex obtuse to rounded, petiole slender
half length of lamina. Inflorescence a rigid, straight, terminal panicle sometimes
reduced to a single axis. Flowers sessile, densely mealy outside, perianth 5-partite.
Male flowers: perianth globular, 0.5-1 mm high, filaments united at base into a thick
broad saucer-shaped disc, glabrous; pistillode hemispherical to obturbinate c. 0.5 cm
high with a pair of minute erect subulate stigmas (rarely the pistillode is fertile). Fe-
male flowers: perianth at first soft and globular to broadly turbinate c. 1 mm high,
eventually spreading under the fruit (never enveloping it) and 2.5-3 mm diameter,
becoming thick and hard and often pale red; staminodes lanceolate, c. 0.3 mm long,
with minute antherodes, united at base into a broad glabrous or sparsely hairy disc;
ovary with a pair of spreading stigmas. Berry depressed globular, pericarp fleshy,
pink or red. Seed lenticular c. 1 mm diameter; testa black, prominently reticulate
with a radially elongated matrix.

QUEENSLAND: Noondoo, S. T. Blake 10543 (BRI); 20 mi [32.2 km] W of Tara, T.
J. McDonald 71 (BRI).

NEW SOUTH WALES: Woorandara Stn., E. M. Canning 3808 (CBG); 16 mi
[25.7 km] W of Garah, K. L. Soiling 332 (NSW).

VICTORIA: 5 mi [8 km] NNW of Kerang, H. J. Aston 559 (MEL); 10 mi [16.1km]
W of Cohuna, J. Vickery 2021 (NSW).

SOUTH AUSTRALIA: 6 km W of Hawker, B. Copley 215 (AD): Tallaringa Well, T.
R. N. Lothian 2689 (AD).

WESTERN AUSTRALIA: 3 mi [4.8 km] S of Reid, T. E. H. Aplin 1677 (PERTH);
Nunna, A. W. H. No. E 17 (PERTH).

NORTHERN TERRITORY: Mt Brassey, A. C. Beauglehole 44701 (NT); Hale River
Plot, T. S. Henshall 930 (NT).

Distribution. Central and eastern Australia. Map 34.

Habitat. Extremely variable due to the numerous ecotypes involved.

Paul G. Wilson, Tribe Chenopodieae

231

Lectotypification of the name Rhagodia spinescens var. deltophylla. This name has
been applied in herbaria by F. Mueller and by later botanists to broad leaved vari-
ants of several species, principally R. spinescens, Chenopodium desertorum and C.
curvispicatum. When searching for a suitable lectotype a number of specimens were
found in herbaria K and MEL which had been collected prior to the publication of
R. spinescens var. deltophylla and which Mueller had so annotated (or as R.
deltophylla), and had evidently referred to when drawing up the protologue; how-
ever, none were labelled as having been collected at Lake Campbell, the principal lo-
cality mentioned therein. I have therefore selected as lectotype a specimen (MEL
87960) which agrees with the original description and which is the only collection I
could find of this taxon that was gathered on the expedition by Babbage to the
“North-western interior of South Australia” in 1858. The specimen is labelled
“Scrub N. E. of Lake Gairdner”; since Lake Campbell is about 65 km NE of Lake
Gairdner it is possible that this specimen is the one referred to by Mueller when he
cited that locality. The name “Rhagodia deltophylla” was used by Mueller in his
Second General Report (1854) but without a description. A specimen of R.
spinescens which bears the name R. deltophylla was collected by him along the River
Murray in January 1854 (herb. K); it presumably represents the plant referred to in
the 1854 Report and is also evidently a syntype of R. spinescens var. deltophylla.
Scott (1978a) gave the type citation of that variety as “South Australia, Lake
Campbell; F. Mueller; Jan 1854”; he evidently had in mind the locality cited by
Mueller for the specimen collected on the Babbage Expedition in 1858 and confused
it with the data on the Mueller collection from the Murray River. Those specimens
cited by Scott under R. deltophylla, that I have been able to check, are of R.
spinescens, Chenopodium desertorum and C. curvispicatum. A similar mixture of
species was included by J. M Black in his private herbarium under R. spinescens var.
deltophylla.

Rhagodia spinescens, as delimited in this paper, exhibits considerable variability,
more, in fact, than has been accepted for any other species of Rhagodia. The leaves
vary in shape from narrow elliptic c. 5 mm long to deltoid and 20 mm long, and in
texture from papery to fleshy.

I have not been able to discriminate clearly distinct infraspecific taxa since the dif-
ferent variants appear to grade into each other and the leaves vary in shape and tex-
ture on the same plant. Some variants are rather distinctive; these appear to be very
limited in their distribution and since they also exhibit variability it is not clear as to
whether they should be treated separately. The more easily recognisable of these
variants are as follows;

(1) Glossy variant. Leaves narrow elliptic to deltate, 4-20 mm long, thin or fleshy;
indumentum thin below of irregularly shaped glossy thin-walled bladder hairs,
almost glabrous above. Staminodes glabrous, disc almost absent. North west
Victoria.

Representative collections. VICTORIA: Hattah Lakes N.P., 13 Mar 1969, G.
Anderson (MEL 88273); 10 mi [16.1 km] NNW of Underbool, A. C. Beauglehole
40379 (MEL); Wyperfeld N.P., A. C. Beauglehole 29535 (MEL).

This variant appears to grade into the local small-leaved variant of typical R.
spinescens which is found in north western Victoria and neighbouring areas of
South Australia and New South Wales.

(2) Bisexual variant. Leaves ovate, small, c. 5 mm long. Flowers bisexual, disc
puberulous. Testa faintly rugose. Indumentum more finely furfuraceous than in
typical variant. Northern South Australia.

232 Nuytsia Vol. 4, No. 2 (1983)

Representative collections. SOUTH AUSTRALIA: 36 km W of Abminga siding,
Lazarides 8276 (AD); Lindsay Ck., B. Major 65 (AD).

(3) Central Australian foetid variant. Leaves deltate to hastiform c. 10 mm long,
closely furfuraceous with very small round bladder hairs. Flowers sessile or pedi-
cellate. Testa almost smooth. Simpson Desert.

Representative collections. NORTHERN TERRITORY: Umbeara Stn., T. S.
Henshall 1545 (NT); Beddome Range, T. S. Henshall 1519, 1533 (AD).

Rather variable and possibly intergrading with the typical variant which is
found in the same area. The foetid plant appears to occur on rocky hillsides
whereas the typical variant is found (in this area) in sand.

(4) Thin leaved variant. Leaves broadly elliptic, thin, c. 10 mm long, finely
furfuraceous with very small irregular bladder hairs. Queensland and Northern
Territory.

Representative collections. QUEENSLAND: Laglan Stn., L. S. Smith 10315
(BRI); Taroom, R. W. Johnson 696 (BRI); 10 mi [16.1 km] N of Wandoon, Speck
1959 (CANB). NORTHERN TERRITORY: 28.7 mi [46.2 km] SW of Ooratippra
H.S., G. C. Chippendale (NT 3528).

Recorded as having a nauseous, fishy odour attractive to flies. Speck states
that the creeping stems root at frequent intervals.

The intergradation between the variants listed and those more typical of R.
spinescens is such as to make any formal classification difficult to apply.

Although Rhagodia spinescens as I delineate it is a very variable species, it is not
so variable as has been considered by some authors. A. J. Scott (1978a), for instance,
perceived it more broadly, including under this name specimens which I recognise as
R. drummondii (Wilson 6088b), R. ulicina (Symon 4659), R. eremaea (Latz 114), and
Chenopodium desertorum (Cunningham 384). The name R. prostrata Moq. was also
cited by him as a synonym of R. spinescens whereas it is a synonym of C. des-
ertorum.

Rhagodia spinescens is evidently most closely related to R. drummondii and R.
eremaea. Apart from differences in leaf shape the latter two species may be dis-
tinguished by the absence of disc or staminodes in the female flowers, the inflor-
escence shape and, in R. eremaea, by the minute irregularly shaped vesicular hairs.

10. Rhagodia eremaea Paul G. Wilson, sp. nov. Figure 5C.

Rhagodia sp. B, Paul G. Wilson in Jessop, FI. Central Australia 53 (1981).

Frutex dioecius ad 2 m altus. Folia alterna ovata vel elliptica 0.5-2 cm longa
(petiolo 0.3-1 cm longo incluso) furfuracea cum pilis minutis vesicularibus
irregulariter formatis ornatis vel glabrescentia. Flores congest! in paniculam ter-
minalem pyramidalem vel paniculam gracilem spiciformen continuam dispositi. Flos
masculus disco patelliformis intra pubescent!. Flos femineus staminodiis plerumque
absentibus.

Typus: Rudall River, 22°33'S, 122°11'E; In rocky loam by creek-line. Much-
branched shrub to 1 m, fruit deep red-pink, 19 May 1971, A. S. George 10676 (holo:
PERTH; iso: CANB, K, MEL, NSW).

Rounded dioecious shrub to 2 m high becoming a scrambler in shade. Branches
slender, closely grey-mealy when young with short branched vesicular hairs. Leaves
alternate, ovate to elliptic, 0.5-2 cm long including petiole c. 1/2 length of lamina.

233

Paul G. Wilson, Tribe Chenopodieae

grey mealy with minute irregularly shaped hairs, glabrescent above. Inflorescence a
terminal pyramidal to slender spicoid panicle, flowers congested so as to appear con-
tinuous along slender branches of panicle or (in northern Australia) in slender
spicoid inflorescences with disjunct glomerules. Perianth 5-partite, densely mealy
with short branched vesicular hairs. Male flower caducous; perianth globular, c.
1.5 mm diam.; filaments united into a thick saucer-shaped disc pubescent within;
pistillode minute (c. 0.2 mm high). Female flower: perianth at first soft and globular
c. 1 mm high, eventually hard and spreading beneath fruit becoming red within;
staminodes usually absent (present in terminal flowers); ovary glabrous with a pair
of slender stigmas. Berry depressed globular, c. 3 mm diameter; pericarpfleshy, red.
Seed depressed globular, c. 1.3 mm diam.; testa black with reticulate ornamentation.

SOUTH AUSTRALIA: Ernabella, 9 Aug. 1933, J. B. Cleland (AD); 96 km W of
Mabel Creek H.S. T. R. N. Lothian 3778 (AD).

WESTERN AUSTRALIA: Finucane Is., 5 Feb. 1981, G. Craig (PERTH); Lorna Glen
Stn., S. Hill 25 (PERTH).

NORTHERN TERRITORY: Ayers Rock,G. Chippendale (NT 663); 20 mi [32.2 km]
NW of Alice Spr., D. J. Nelson 1703 (NT).

Distribution. Central Australia westwards. Map 31.

Habitat. Recorded in clay-loam and on rocky hillsides with Acacia kempeana, also on
sand with spinifex.

The flowers are arranged in glomerules each of which is subtended by a caducous
linear bract. These glomerules may consist of a single terminal flower or this flower
may be surrounded by others which may or may not be bracteate. The terminal
flower of a glomerule in a female plant has staminodes while in the lateral flowers
these are usually absent.

This species exhibits the following geographic variation. In central Australia the
leaf is oblong-elliptic with rounded apex and somewhat leathery when dry. In the
north-west of Western Australia (Hamersley Range area) the leaf is narrow-elliptic
with acute apex and is thinner in texture. The plant from the north-west and from
the far north of both Western Australia and the Northern Territory usually has a
very slender, spicate type of inflorescence with discontinuous glomerules of flowers,
unlike the continuous, dense spikes of the central Australian plant. Another variarit
found in Western Australia between Geraldton and Wittenoom has thin, broadly el-
liptic to obovate leaves with rounded apices; the leaves of those specimens growing in
shade are thinner and broader than those in the open.

In herbaria female plants of R. eremaea have generally been identified as R.
spinescens (or, more rarely, as R. crassifolia) and male plants as Chenopodium
nitrariaceum. From both R. spinescens and R. crassifolia this species may be dis-
tinguished by the very small irregularly shaped vesicular hairs of the leaf indumen-
tum, the slender, compact inflorescence, and by the lack of staminodes in the remale
flowers. Male plants may be distinguished from C. nitrariaceum by the indumentum,
the scattered leaf arrangement, and the absence of spinescent branchlets.

A specimen of R. eremaea collected by F. Mueller in 1856 along Sturt Creek
(north-east Western Australia) is evidently one of several syntypes of Chenopodium
nitrariaceum (F. Muell.) Benth. This specimen was later re-identified by Bentham
(1870) as R. crassifolia which name was also applied by Ewart and Davies (1917).

Nuytsia Vol. 4, No. 2 (1983)

The specific epithet is taken from the Greek word eremia, a desert, alluding to the
typical habitat of this plant.

11. Rhagodia parabolica R. Br., Prod. 408 (1810). Type: Inlet XII [Spencers
Gulf], March 1802, R. Brown (holo: BM).

R. parabolica var. typica Domin, Bibl. Bot. 89:62 (1921) nom. illeg.

R. reclinata A. Cunn. ex Moq. in DC., Prod 13/2:51 (1849).--R. parabolica var
reclinata (Moq.) Domin, Bibl. Bot. 89:62 (1921). Type: Liverpool Plains, May 1825,
A. Cunningham 7 (holo: K).

Rounded dense dioecious shrub 1-2 m high. Branches striate, somewhat flattened
when young, mealy grey with minute irregularly shaped vesicular hairs. Leaves pre-
dominantly opposite or subopposite, chartaceous, broadly or very broadly elliptic to
broadly or very broadly ovate (rarely bluntly hastate) 1.5-4 cm long, apex acute to
rounded, apiculate, base truncate to cuneate passing into the slender petiole half the
length of or equal in length to lamina, minutely mealy becoming reddish brown with
age. Inflorescence a broad pyramidal terminal panicle 5-8 cm long, the flowers in dis-
crete glomerules along its branches. Flowers spherical c. 1 mm diameter; perianth 5-
partite, closely mealy, eventually spreading in the female flower to 2.5 mm diam. but
scarcely indurated. Male flowers: staminal filaments united at base into a thick cu-
pular disc which is densely wooly at junction with filaments; pistillode minute. Fe-
male flowers: staminodes in terminal flower of glomerule prominent, linear with min-
ute antherodes and united into a deep cupular disc densely wooly within, in lateral
flowers reduced or absent; stigmas subulate and spreading. Berry depressed globular
c. 2.5 mm diameter. Seed depressed globular to lenticular, c. 1.8 mm diam. testa
glossy with faint radial ribbing.

QUEENSLAND: Online, S. T. Blake 10930 (BRI); Isle Gorge, P. Sharpe 00646

WALES: Mt Hope, G. M. Cunningham and P. Milthorpe 2511
(NSW); 6 mi [9.7 km] W by S of Jerrys Plains, R. Story 7131 (CANB).

VICTORIA: Werribee Gorge, A. C. Beauglehole 39927 (MEL); Head of Coimadai
Creek, 7 May 1944, J. H. Willis (MEL).

Koondoolka, J. Z. Weber

N()RTHERN TERRITORY; Ilpapa Swamp, T. S. Henshall 1196 (NT);
[0.8 km] W of Ooraminna Rockhole, D. J. Nelson 1964 (MEL).

0.5 mi

Distribution. South-east Queensland and north east New South Wales, southern Vic-
toria, southern South Australia and Central Australia. Map 33.

2nd banks^'''’'''*''^ hillsides with skeletal soil, along creek banks and on

This species appears to occupy four disjunct areas in Australia but the plants from
each area are sufficiently alike not to warrant infraspecific recognition.

In the Northern Territory occasional plants are hermaphrodite and some have
tical (not horizontal) seeds, but it is not possible to establish from herbarium
tenal now common these variants are.

ver-

ma-

Paul G. Wilson, Tribe Chenopodieae

235

In Yorke and Eyre Peninsulas and north of Adelaide, plants are found with elliptic
leaves and a hair type intermediate between those found in typical R. parabolica
(with small discrete vesicular hairs) and that found in R. candolleana (in which the
hairs collapse to form a scurfy or silvery sheen). The typical, broad-leaved R. para-
bolica has been collected in the same areas so that it is possible that the narrow-
leaved plant has arisen through hybridization with another species such as R. preissii
or R. candolleana.

Doubtful or incorrect records

Blituw virgatum L. (= Chenopodium foliosum (Moench) Asch.) Recorded by J. D. Hooker, FI. Tasman.
XCVI (1959) as being found in Australia. I have seen no specimens which support this record.

Chenopodium bonus-henricus L. In the Victorian Naturalist 22:79 (1905) it is recorded that H. B.
Williamson exhibited a specimen of this species collected from ‘Hopkins mouth’, Victoria. This record
is referred to by Ewart and Tovey (1909) and by Ewart (1931) who state that it was naturalized in
south-west Victoria in 1905. 1 have been unable to locate the Hopkins mouth collection but a specimen
collected by Williamson nearby at Hawksdale in 1900 (MEL 88604 and 88605), and identified by
Ewart as C. bonas-henricus, is of a species of Atriplex.

Chenopodium iicifolium Smith. Recorded as a weed in Brisbane by F. M. Bailey, Queensland Agric. J. 31:
321 (1913). I have seen no specimens to validate this record.

Chenopodium rubrum L. Recorded by Audas and Morris (1925) as widely spread in Victoria and by Ewart
(1931) as naturalized at North Melbourne in 1910 and widely spread in Victoria. I have seen no Aus-
tralian collection of this species. The following two specimens in herb. MEL which had been deter-
mined as C. rubrum are, in fact, both Chenopodium glaucum: Railway Reserve, N. Melbourne, May
1910, C. French and R. J. Tovey (MEL 88793); Coode Island, 23 March 1912, R. J. Tovey (MEL
88794).

Chenopodium viride L. Recorded by J. D. Hooker, FI. Tasman. CIX (1859) as being found in Australia. I
have seen no specimen to support this record.

Excluded names

Chenopodium insulare J. Black, Trans. Roy. Soc. S. Austral. 69: 309 (1945) = Suaeda australis (R. Br.)
Moq. See Hj. Eichler, Supplement to J. M. Black’s Flora of South Australia (1965).

Chenopodium australe R. Br,, Prod. 407 (1810) = Suaeda australis (R. Br.) Moq.

Rhagodia eschscholtziana Fenzl, Denkschr. Akad, Wien (Math. -Nat.) 1: 257 (1850). Described from Oahu
in the Hawaiian Islands; the description agrees with that for Chenopodium oahuense (Meyen) Aellen
(Atriplex oahuensis Meyen 1835).

Acknowledgements

I am grateful to the Directors of the State, Commonwealth, and University
herbaria in Australia and to the herbarium of the DSIR in Christchurch, for the loan
of material. I am also grateful to the Directors of the herbaria in Paris (P), Florence
(FI), Lund (LD), London (BM), and Kew (K) for the loan of type specimens. The
National Herbarium of New South Wales (NSW) kindly sent manuscript herbarium
notes on many of the species treated in this paper. Mr W. R. Sykes (CHR) sent com-
ments on several of the species found in New Zealand.

The present study was commenced in 1978 while a guest of the Division of Plant
Industry, CSIRO, Canberra. I am grateful to its Chief, Dr W. J. Peacock, for ex-
tending an invitation and to Dr Hj. Eichler, then Curator of the Herbarium Aus-
traliense, for making space and facilities available and for his personal support. The
SEM photographs, figures 4 and 5, were prepared by Mr Stuart Craig of the Division
of Plant Industry, Canberra and figures 6 and 7 by Mr Terry Robertson of the Queen
Elizabeth II Medical Centre, Perth, to both of whom I wish to express my appreci-
ation for their careful work.

236

Nuytsia Vol. 4, No. 2 (1983)

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Giusti, L. (1970). El genero Chenopodium en Argentina. 1. Numeros de cromosomas. Darwiniana 16: 98-
105.

Hellmuth, E. O. (1968). Eco-physiological studies on plants in arid and semi-arid regions in Western Aus-
tralia. 1. Autecology of Rhagodia baccata (Labill.) Moq. J. Ecol. 56: 319-344.

Hooker, J. D. (1880). Chenopodiaceae. In G. Bentham and J. D. Hooker, ‘Genera plantarum’ vol. 3. (Reeve:
London.)

Jorgensen, P. M. (1973). The genus Chenopodium in Norway. Norw. J. Bot. 20: 303-319.

Kjellmark, S. (1934). Einige neue Chromosomenzahlen in der Familie Chenopodiaceae. Bot. Not. 136-149.

Maiden, J. H. and Betche, E. (1916). ‘A census of New South Wales Plants.’ (Government Printer:
Sydney.)

Moquin-Tandon, C.H.B.A. (1840). “Chenopodearum monographica enumeratio.” (P. J. Loss: Paris.)

Moquin-Tandon, C.H.B.A. (1849). Salsolaceae. In A. P. de Candolle, “Prodromus systematis naturalis
regni vegetabilis” vol. 13/2. (Teuttel et Wiirtz: Paris.)

Mueller, F. (1859). “Report on the plants collected during Mr. Babbage’s expedition into the north-west
interior of South Australia in 1858.” (Government Printer: Melbourne.)

Mueller, F. (1869-1871). “Fragments Phytographiae Australiae” vol. 7. (Government Printer: Melbourne.)
Mueller, F. (1886). New Australian plants. Australas. J. Pharm. 1:429-431.

Mueller, F. (1887). New Australian Plants. Bot. Centralbl. 29:114-115.

Mueller, F. (1889-1891). “Iconography of Australian salsolaceous plants.” (Government Printer:
Melbourne.)

Murr, J. (1910). Australische Chenopodien. Allg. Bot. Zeit. 16:55-58.

Murr, J. (1926). Meine Phanerogamen-Bastarde. Vierteliahrsschr. Gesch. Landesk. Voralbergs 10:185-194.
194.

Murr, J. and Thellung, H. (1915). Chenopodium auricomiforme. Vierteliahrsschr. Naturf. Ges. Zurich
60:432.

Osmond, C. B. et al. (1969). Ion absorption in Atriplex leaf tissue. II. Secretion of ions to epidermal blad-
ders. J. Biol. Sci. 22:797-814.

Pax, F. A. (1889). Dysphania. In A. Engler and K. Prantl, “Die natiirlichen Pflanzen-familien” 3, lb:92.
(Verlag von Wilhelm Englemann:Leipzig.)

Pax, F. A. (1927). Zur Phylogenie der Caryophyllaceae. Bot. Jahrb. Syst. 61:223-241.

Pax, F. A. and Hoffmann, K. (1934). Dysphaniaceae. In A. Engler and K. Prantl, “Die naturlichen
pflanzenfamilien” ed. 2, 16c: 272-274. (Duncker and Humblot: Berlin.)

Planchuelo, A. M. (1975). Estudio de los frutos y semillas del genero Chenopodium en la Argentina.
Darwiniana 19:528-565,

Probst, R. (1949). “Wolladventiflora Mitteleuropas.” (Vogt-Schild Ag.: Solothurn.)

Rodway, L. (1903). “The Tasmanian flora.” (Government Printer: Hobart.)

Scott, A. J. (1978a). Rhagodiinae: A new subtribe in the Chenopodiaceae. Feddes Repert. 89:1-11.

238 Nuytsia Vol. 4, No. 2 (1983)

Scott, A. J. (1978b). A review of the classification of Chenopodium L. and related genera
(Chenopodiaceae). Bot. Jahrb. Syst. 100:205-220.

Shapter, R. E. (1941). A preliminary investigation of the yield and composition of the oil distilled from
Chenopodium ambrosioides (Linn.) var. anthelminticum (Gray). J. Council Sci. Ind. Rea. 14:201-208.

Standley, P. C. (1931). The Chenopodiaceae of northwestern South America. Field Mus. Nat. Hist. Bot
11:115-126.

Sykes, W. R. 11977). "Kermadec Islands Flora.” Bulletin no. 219. (New Zealand D.S.I.R.: Wellington.)

Ulbrich, E. (1934). Chenopodiaceae. In Engler et Prantl, “Die natiirlichen Pflanzenfamilien” ed. 2, 16c:379-
584. (Duncker and Humblot: Berlin.)

Uotila, P. (1978). Variation, distribution and taxonomy of Chenopodium suecicum and C. album in N.
Europe. Acta Bot. Fennica 108:1-35.

Voroschilov, V. N. (1942). Revision des especes de Chenopodium de la section Ambrina (Spach) Hook.f. J.
Bot. URSS 27 No. 3/4:33-47.

White, C. T. (1921). Illustrated notes on the weeds of Queensland. No. 21. Goosefoot (Chenopodium
triangulare, R.Br.). Queensld. Agric. J. n.s. 15:171-172.

Willis, J. H. (1973). “A handbook to plants in Victoria.” vol. 2 (Melbourne Univ. Press: Victoria.)

Wilson, H. D. (1980). Artificial hybridization among species of Chenopodium sect. Chenopodium Svst
Bot. 5:253-263.

Wood, J. G. (1923). On transpiration in the field of some plants from the arid portions of South Australia
with notes on their physiological anatomy. Trans. Roy. Soc. S. Austral. 47:259-278.

Paul G. Wilson, Tribe Chenopodieae

239

Figure 4. SEM photos of seeds: A — Chenopodium erosum (upper surface with pericarp attached), from
A. C. Beauglehole 37201 (PERTH); B — Chenopodium murale (lower surface), from CANB 31776;

C — Chenopodium auricomiforme (lower surface), from CANB 36430; D — Chenopodium curvispicatum
(lower surface), from H. Schodde 1082 (CANB); E — Einadia nutans subsp. nutans (lower surface),
from AD95826008; V- Einadia hastala (lower surface), from CBG04660t.

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Figure 5. SEM photos of seeds: A — Einadia allanii (upper surface), from P. Waidle (CHR125728);
B — Einadia triandra (upper surface), from iV. J. A. Simpson 7160 (CHR); C — Rhagodia eremaea
(lower surface), from P. K. Latz 139 (PERTH); D — Rhagodia spinescens (upper surface), from
ADW34053; E — Rhagodia acicularis (upper surface), from P. G. Wilson 11713 (PERTH);
F — Rhagodia ulicina (upper surface), from AD96503078.

Paul G, Wilson, Tribe Chenopodieae

241

Figure 6. SEM photos of seeds: A—Dysphania glomulifera subsp. glomulifera (side view), from
C. Andrews, April 1902 (PERTH); B—Dysphania glomulifera subsp. eremaea (side view showing em-
bryo-groove), from R. D. Royce 1493 (PERTH); C^Dysphania glandulosa (side view showing embryo-
groove), from A. S. George 717 (PERTH); D—Dysphania platycarpa (side view), from A. S. George
8927 (PERTH); E-F—Dysphania sphaerosperma (side view showing vertical embryo-groove).

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Figure 7. SEM photos of seeds; A — Dysphania kalpari (side view showing raphe), from D. J. Nekon
830 (NT); B — Dysphania littoralis (obliquely lateral view), from P. G. Wilson, 24 Feb. 1977 (PERTH);
C-D — Dysphania rhadinostachya (C — obliquely lateral view and D — upper surface, showing horizontal
embryo-ridge), from R. J. Ch/niioclf 385,1 (PERTH); E — Dysphania plan tagfne/la (side view), from A.
C. Beauglehole 48362 (PERTH); F — Chenopodium gaudichaudianum (upper surface), from P. G.
Wilson 9911 (PERTH).

Paul G. Wilson, Tribe Chenopodieae

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Nuytsia Vol. 4, No. 2 (1983)

Index to scientific names

Numerals in bold type represent main entries of accepted names.

New names are given in bold type. Other accepted names are in Roman. Synonyms
are in italics.

Amaranthus L. 197

enervis (F. Muell.) F. Muell. 197
tenuis Benth. 197

Ambrina Spach 166

sect. Adenois Moq. 166
sect. Botryois Moq. 180
carinata (R. Br.) Moq. 173
var. parvifolia Moq. 174
pinnatisecta Spach 168
punailio (R. Br.) Moq. 172

Atriplex L. 235

glomuHfera Nees 183
oahuensis Meyen 235
vesicaria Benth.

Blitum L.

sect. Orthosporum (R. Br.) C. Meyer 169
Atriplicinum F. Muell. 197
caiinatum (R. Br.) Steudel 173
cristatum F. Muell. 176
glandulosum Moq. 172

var. parvi folium (Moq.) Moq. 174
pumilio (R. Br.) Steudel 172
virgatum L. 235

Chenopodium L. 139

sect. Ambrina J. D. Hook. 169

sect. Ambrina (Spach) J. D. Hook. 166

subg. Ambrosia A. J. Scott 166

sect. Atriplicina Aellen 197

sect. Auricoma Aellen 161

sect. Botryoides C. Meyer 180

sect. Boti^'ois (Moq.) Moq. 180

sect. Carinata Standley 169

sect. Chenopodiastrum Moq. 144

sect. Chenopodium 134

subg. Chenopodium 139

sect. Degenia Aellen 143

sect. Desertorum Paul G. Wilson 151

sect. Dysphania (R. Br.) Aellen 181

sect. Euchenopodium Ulbrich 144

subsect. Lejosperma Aellen et Iljin 144

sect. Leprophyllum Dumort. 144

sect. Margaritaria Brenan 180

sect. Orthosporum R. Br. 139, 169

sect. Polygonoidea Aellen 199

sect. Pseudoblitum J. D. Hook. 141

sect. Rhagodioides Benth. 164

sect. Roubieva (Moq.) Asch. et Graebner 166

sect. Tetrasepala Aellen 181

sect. Thullingia Aellen 197

acuminatum Willd. 164

album L. 144

subsp. opulifolium (Koch et Ziz) Maire 145
var. reticulatum (Aellen) Uotila 145
aWanij Aellen 210
ambiguum R. Br. 141
var. majus Moq. 141
var. minus Moq. 141
ambrosioides L. 167
var. anthelminticum (L.) A. Gray 167
f. spathulatum (Moq.) Aellen 167
anidiophyllum Aellen 152, 155
antarcticum (J. D. Hook.) J. D. Hook. 198
anthelminticum L. 167
atriplicinum (F. Muell.) F. Muell. 197
auricomiforme Murr et Thellung 162
auricomum Lindl. 163
f. subglabrum Aellen 163
australasicum Moq. 200
australe R. Br. 235
baccatum Labill. 223
berlandieri Moq. 146
bi forme Nees 148
blackianum Aellen 188
xbontei Aellen 171
n-var. cristatiforme Aellen 171
n-var. submelanocarpum Aellen 171
botrys L. 180
bonus-henricus L. 235
browneanum Roemer et Schultes 144
carinatum R. Br. 173
f. foliosum Domin 174
var. holopterum (Thellung) Aellen 174
f. leucocarpum Aellen 176
var. melanocarpum J. Black 175
f. rubellum Aellen 172
carnosulum Moq. 148
xchristii Aellen 171

n-var. intermedium Aellen 171, 172
n-var. semiconnatum Aellen 171, 172
cochlearifolium Aellen 152
congestum J. D. Hook. 148
congolanum (Haum.) Brenan 180
cristatum (F. Muell.) F. Muell. 176
var. holopterum Thellung 174
curvispicatum Paul G. Wilson 159
desertorum (J. Black) J. Black 152

subsp. anidiophyllum (Aellen) Paul G.

Wilson 155

subsp. desertorum 153
subsp. microphyllura Paul G. Wilson 154
subsp. rectum Paul G. Wilson 158
subsp. virosum Paul G. Wilson 157

Paul G. Wilson, Tribe Chenopodieae

detestans J. Kirk 147
erosum R. Br. ISO
ficifolium Smith 235
foliosum (Moench) Asch. 235
furfuraceum Moq. 214

gaudichaudianum (Moq.) Paul G. Wilson 160
giganteum D. Don 145
glandulosum (Moq.) F. Muell. 172
glaucum L. 139

var. ambiguum (R.Br.) J. D. Hook. 141
subsp. ambiguum (R.Br.) Thell. 141
var. littorale Rodway 141
f. minus (Moq.) Aellen 141
var. paschale Fuentes 141
holocarpum J. Black 175
hoJopterum (Thell.) Thell. et Aellen 174
hubbardii Aellen 151
inflatum Aellen 193, 194
insulare J. Black 235
integrifolium Vorosch. 168
lanceolatum R.Br. 144
littorale (L.) Thunb. 141, 188
littorale Moq. 141
lycioides Gand. 164
macrospermum J. D. Hook. 143
subsp. halophilum (Philippi) Aellen 143
f. subviride Aellen 143
melanocarpum (J. Black) J. Black 175
f. leucocarpura (Aellen) Paul G. Wilson 176
f. melanocarpum 175
microphyllum F. Muell. 152, 154
var. desertorum J. Black 152
microphyllum Thunb. 152
multifidum L. 168
f. intermedium Aellen 168
f. spathulatum Aellen 168
murale 148

var. bi forme (Nees) Moq. 148
var. sinuosifrons Murr 148
myriocephalum (Benth.) Aellen 183
nitrariaceum (F. Muell.) Benth. 164
oahuense (Meyen) Aellen 235
olidum Curtis 146
opulifolium Koch et Ziz 145
osbornianum Aellen 192
plantnginellum (F. Muell.) Aellen 188
polygonoides (Murr) Aellen 212
preissii (Moq.) Diels 221
probstii Aellen 144
f. lanceolatum Aellen 144
f. parvo-angustifolium Aellen 144
f. subintegrifolium Probst 144
prostratum (Pallas) Roemer et Schultes 139
pseudomicrophyllum Aellen 152, 154
pumilio R.Br. 172
f. glandulosum (Moq.) Aellen 172
var. oblongifolium J. Black 172
f. rubellum (Aellen) Aellen
pussillum J. D. Hook. 179
22493 — ( 9 )

261

reticulatum Aellen 144
rhadinostachyum F. Muell. 193
var. paniculatum F. Muell. et Tate 193
rubrum L. 142, 235
saxatile Paul G. Wilson 179
scabricaule Speg. 148
serotinum L. 145
simulans (Tate) F. Muell. 192
spathulatum Sieberex Moq. 167, 168
stellulatum Aellen 206, 208
stellulatum (Benth.) Aellen 208
suecicum Murr 145, 150
suffruticosum Willd. 168
subsp. remotum Vorosch. 168
tenue Colla 139
triandrum G. Forster 205
var. lanuginosum F. E. Haviland 152, 154
triangulare Forsskal 145
triangulare R.Br. 206
var. angustifolium Benth. 212
subsp. convohulinum Murr 202
var. convolvulinum (Murr) Maiden 202
subsp. polygonoides (Murr) Murr 212
var. polygonoides Murr 212
var. stellulatum Benth. 208
trigonocarpum Aellen 171
trigonon Roemer et Schultes 206
truncatum Paul G. Wilson 177
ulicinum Gand. 219
viride L. 235
vulvaria L. 146

Dysphania R.Br. 180

sect. Caudatae A. J. Scott 181
sect. Tetrasepalae (Aellen) A. J. Scott 181
benthamiana Domin 183, 184
glandulosa Paul G. Wilson 186
glomulifera (Nees) Paul G. Wilson 183
subsp. glomulifera 184
subsp. eremaea Paul G. Wilson 184
inf lata (Aellen) A. J. Scott 193
kalpari Paul G. Wilson 195
littoralis R.Br. 188
myriocephala Benth. 183
plantaginella F. Muell. 190
platycarpa Paul G. Wilson 189
rhadinostachya (F. Muell.) A. J. Scott 193
subsp. inflata (Aellen) Paul G. Wilson 194
subsp. rhadinostachya 194
simulans Tate 192
sphaerosperma Paul G. Wilson 191
valida Paul G. Wilson 187

Einadia Raf. 199

allanii (Aellen) Paul G. Wilson 210
hastata (R.Br.) A. J. Scott 211
linifolia (R.Br.) Raf. 200, 204
nutans (R.Br.) A. J. Scott 200
subsp. eremaea Paul G. Wilson 204
subsp. linifolia (R.Br.) Paul G. Wilson 204

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Nuytsia Vol. 4, No. 2 (1983)

var. lini folia (R.Br.) A. J. Scott 204
subsp. nutans 202

subsp. oxycarpa (Gauba) Paul G. Wilson
203

polygonoides (Murr) Paul G. Wilson 212
triandra (G. Forster) A. J. Scott 205
trigones (Roemer et Schultes) Paul G. Wilson
206

subsp. leiocarpa Paul G. Wilson 209
subsp. stellulata (Benth.) Paul G. Wilson
208

subsp. trigonos 209

Euxolus Raf.

enervis F. Muell. 197

Monolepis Schrader 181, 197, 199

Orthospermum (R.Br.) Opiz 169

Orthosporum (R.Br.) T. Nees 169
bonus -henriciis (L.) T. Nees
rubrum (L.) T. Nees

Rhagodia R.Br. 213

acicularis Paul G. Wilson 221
baccata (Labill.) Moq. 223
var. angustifolia Moq. 224
subsp. baccata 223.
var. candolleana (Moq.) Moq. 214
var. congesta (J. D. Hook.) J. D. Hook. 148
subsp. dioica (Nees) Paul G. Wilson 225
var. linearis (Benth.) J. Black 223
var. parx'ifolia Moq. 215
billardieri R.Br. 223
var. congesta (J. D. Hook) Benth. 148
var. linearis Benth. 223
candolleana Moq. 214
subsp. argentea Paul G. Wilson 215
subsp. candolleana 215
cbenopodioides Moq. 200, 202
congesta (J. D. Hook.) Moq. 148
coralliocarpa S. Moore 160
crassifolia R.Br. 218
var. latifolia Benth. 228
cygnoruni Gand. 223
deltophylla (F. Muell.) A. J. Scott
deltophylla F. Muell.

dioica Nees 223, 225
drummondii Moq. 229
eremaea Paul G. Wilson 232
eschscholtziana Fenzl 235
gaudichaudiana Moq. 160
hastata R.Br. 211

latifolia (Benth.) Paul G. Wilson 228
subsp. latifolia 228
subsp. recta Paul G. Wilson 228
linifolia R.Br. 200, 204
nitrariacea F. Muell. 164
nutans R.Br. 200
var. fallacina Domin 202
var. oxycarpa Gauba 202
var. parvifolia Moq. 202
obovata Moq. 221, 222
parabolica R.Br. 234
var. reclinata (Moq.) Domin 234
var. typica Domin 234
parvifolia Moq. 218
preissii Moq. 221

subsp. obovata (Moq.) Paul G. Wilson 222
subsp. preissii 222
prostrata Moq. 152
radiata Nees 223
reclinata Nees 234
spinescens R.Br. 230
var. deltophylla F. Muell. 230
triandra (G. Forster) Aellen 205
ulicina (Gand.) Paul G. Wilson 219

Roubieva Moq. 166

multifida (L.) Moq. 168

Salsola L.

carinata (R.Br.) Sprengel 173

Scleroblitum Ulbrich 197

atriplicinum (F. Muell.) Ulbrich 197

Suaeda Forsskal ex Scop. 199
australis (R.Br) Moq. 235
linifolia Pallas 200

Tetragonia L.

iraplexicoma (Miq.) J. D. Hook. 218

Publication date of Nuytsia Volume 4 Number 1: 14 Oct. 1982.

22493/1 1 /82— 1 M— L/3286

WILLIAM C BROWN. Government Printer. Western Australia

Notes for Authors

Nuytsia publishes papers relating to the flora of Western Australia. All papers are
refereed outside the Western Australian Herbarium. The Herbarium reserves the
right to reject papers.

Manuscripts must be submitted in duplicate. They should be typewritten, double
spaced, with 2.5 cm margins and no underlining. Pages should be numbered. The de-
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provided. For style and layout, recent numbers of Nuytsia should be followed. Galley
proofs will be forwarded to authors for checking. Reprints may be ordered at the
author’s expense.

CONTENTS

A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in

Australia. By Paul G. Wilson 135

Publication date of Volume 4 Number 1 262